Handbooks for the Identification of l3rltmh lnseets Va1. 10, Part 6

SCUTTLE FLIES DIPTERA, PHORIDAE

(except Megase/ia)

R.H.L. Disney

Handbooks for the Identification of British Insects

Editor: M.G. Fitton

SCUTTLE FLIES DIPTERA, PHORIDAE

(except Megaselia)

By

R.H.L. Disney Malham Tarn Field Centre

Settle North Yorkshire BD24 9PU

1983

VoL 10, Part 6

ROYAL ENTOMOLOGICAL SOCIETY OF LONDON

The aim of the Handbooks is to provide illustrated identification keys to the insects of Britain, together with concise morphological, biological and distributional information. The series also includes a Check List of British Insects.

Each handbook should serve both as an introduction to a particular group of insects and as an identification manual.

Details of handbooks currently available, and an order form, can be obtained from the Royal Entomological Society, 41 Queen's Gate, London SW7 5HU.

World List abbreviation: Handbk Ident. Br. Insects.

© Royal Entomological Society of London, 1983

First published 1983 by the Royal Entomological Society of London, 41 Queen's Gate, London SW7 SHU. .

Printed by Dramrite Printers Limited, Southwark, London SEl

Contents

Introduction Relationships with other families. Natural history . Collecting. Preservation and examination Morphology Notes on the Keys. Acknowledgements Check list. Key to genera Genus Aenigmatias Genus Anevrina . Genus Beckerina . Genus Borophaga. Genus Chaetopleurophora. Genus Chonocephalus Genus Conicera . Genus Diplonevra. Genus Dohrniphora Genus Gymnophora Genus Gymnoptera Genus Hypocera . Genus Metopina . Genus Phalacrotophora Genus Pbora Genus Plectanocnema Genus Pseudacteon Genus Puliciphora. Genus Spinipbora. Genus Triphleba . Genus Woodiphora References. Plates and figures. Index

Introduction

Page 3 4 5 7 7 9

11 11 12 15 17 18 19 19 20 20 21 22 23 24 25 25 25 28 28 30 30 31 31 32 37 37 40 79

The scuttle flies (the name refers to their characteristic scuttling gait) or Phoridae are one of the largest families of Diptera. Approximately 300 species occur in the British Isles, of which more than 200 belong to the giant genus Megaselia. This volume covers the 93 species that belong to genera other than Megaselia. These genera embrace some of the largest scuttle flies (up to 8mm in length) as well as some of the smallest (less than Imm in length). The larval natural histories (where known) range from true parasites, through parasitoids, specialised predators, fungus feeders, to polyphagous saprophages. However the biology of most species is unknown. The neglect of this family is a direct consequence of a lack of identification keys which can be used with confidence by the non-specialist. This Handbook aims to remedy this deficiency for the British fauna.

3

Megaselia sea/oris

Relationships with other families

The Phoridae is usually regarded as a distinctive family of the Division Aschiza of the Suborder Cyclorrhapha. being placed in the superfamily Phoroidea along with the Platypezidae, Ironomyiidae and Sciadoceridae. McAlpine & Martin (1966) have discussed the relationship of these families. Subsequently Hennig (1971) reconsidered their assignment of two interesting Cretaceous fossils, transferring the latter from the Sciadoceridae to the Phoridae. However, the opinion of the former authors has much to commend it when attention is concentrated on features other than the simplified wing venataions of these flies. The placing of the Platypezidae in the same group as the rest of the Phoroidea is based on the most tenuous evidence. The views of Griffiths (1972) would seem more soundly based. He places the Phoridae, Ironomyiidae and Sciadoceridae in the group Hypocera. The Platypezidae are removed to a separate group - the Platypezidea. He suggests only further research can settle the question as to

4

whether the latter group are closer to the Hypocera or to the Syrphidea and Schizophora. I am of the opinion that the Hypocera are more sharply demarcated from the rest of the 'Ascruza' than the above review of current opinions suggests ..

The evidence for associating the Phoridae, Ironomyiidae and Sciadoceridae seems strong. The evidence of the wing venation suggests a series running from the Ironomyiid condition, through the Sciadocerid condition, to the Phorid condition. In other respects the first two families seem to possess a range of characters that appear to be apomorphous (derived) with respect to the situation thought to characterise the primitive Phoridae. It has been suggested (Disney, 1981d) that the median furrow of the frons, present in many Phoridae, is a plesiomorphous (ancestral) feature representing the anterior part of the epicranial suture found in many Nematocera. There also appears to be a vestigial median furrow in one Sciadocerid genus (Archiphora) according to the illustration given by Schmitz (1929). Another character thoughtto represent the plesiomorphous state in Phoridae is the two-jointed palp found in some genera (e.g.Dohrniphora).

In most Phoridae the eclosion of the adult from the puparium seems to be initiated by the longitudinal split dividing the eclosion plate into two halves (fig. 5), to be followed by the detachment of the rest of the plate margins. Besides the whole plate is dorsal and does not involve the anterior segments at all. The puparium of Sciadocera is very similar (Fuller, 1934).

It is the sort of considerations reviewed above that tends to revive interest in the view of earlier morphologists (e.g. Wesche, 1908) that the aff'mities of the Phoridae lie more with the Brachycera than with the rest of the Cyclorrhapha. Venturi (1966) considered certain features of the male hypopygium in the Phoridae to be more reminiscent of the Brachycera than the rest of the Cyclorrhapha, thus supporting these earlier views.

It is evident that we need a much clearer idea of which character states encountered in the Phoridae are plesiomorphous and which are apomorphous. While the evidence strongly suggests some affinity between Phoridae and Sciadoceridae and Ironomyiidae the possibility that this group of families represent a line from the Brachycera that evolved independently of that followed by the rest of the Aschiza cannot be ruled out. On the contrary it is a hypothesis that merits serious consideration. Furthermore certain features reminiscent of the Nematocera suggest separation from the Brachycera at an early stage in the evolution of the latter.

Natural History

The typical life history includes an egg, three larval instars and a pupa (enclosed in a 'puparium'). In some tropical species there is known to be suppression of some or all of these stages as free-living entities. The natural histories of the larvae are extremely diverse and what little we know suggests many surprises await to be revealed.

The EGG may be simple as in Megaselia halterata (fig.I), whose larvae feed on the mycelium of mushrooms, or possess longitudinal ridges as in M. rnficornis (fig.2), whose larvae develop in dead snails and other carrion.

The LARVA is characterised by the presence of tapered processes on each body segment (figs. 3-4), recalling the larvae ofPlatypezidae or Fanniidae. These processes are reduced in parasitic species.

The PUP ARIUM (fig. 5) is characterised by the presence of respiratory horns. These horns may be absent in specimens parasitized by Hymenoptera. Eclosion of the adult takes place by detachment of 1;wo dorsal plates from the thoracic region, each plate bearing one of the respiratory horns. Occasionally the two plates may be fused, into a single plate, along the median line. This is the case in Megaseliajuscinervis, whose puparium is distorted to fit the shell of its snail host, the single eclosion plate serving

to occlude the aperture of the shell (Disney, 1982c). Returning to consideration of larval natural histories a few species are polyphagous

saprophages. Megaselia sealaris is, perhaps, the supreme example of this habit in the animal kingdom (see McCrae, 1962, Robinson, 1971). Some species are selective saprophages, for example the species of Spiniphora seem largely to restrict themselves to breeding in dead snails; although S. bergenstammi puparia have been frequently reported from empty milk bottles (e.g. Oldroyd, 1964). A number of species breed in fungi. While some of these (e.g. Megaseiia halterata andM. nigra) are known to feed on fungus tissue others may be predators or parasitoids of other fungivorous insect larvae or their eggs.

The transition to a predatory habit is foreshadowed in the habits of Dohrniphora eornuta. The larvae of this species (fig.3) live in compost heaps and trickling filter sewage beds. In the latter situation they normally feed on the film of micro-organisms coating the substratum. If, however, competition from the larvae of Psyehoda alternata becomes too severe they proceed to prey upon the eggs, larvae and pupae of the latter species (Kloter et al., 1977).

Perhaps the majority of larval Phoridae are specialised predators or parasitoids. Some are true parasites.

Megase[ia aequalis and M. ciliata larvae prey upon slug eggs (Robinson & Foote, 1968, Disney, 1977, 1979b). M .. melanoeephala, M. nasoni and M. pu/iearia larvae feed on spider eggs (Decou-Burghele, 1961, Disney & Evans, 1980). M. rujalarvae prey on coccoids and their eggs (Schmutterer, 1952), While Phora holoserieea larvae prey on root aphids (yarkulov, 1972). M. juscinervis larvae attack Vitraea snails (Disney, 1982c). M. e/ongata larvae parasitize millipedes (Picard, 1930), M. flavieoxa and M. obscuripennis parasitize larval Sciaridae (Disney, 1976, 1980c). M. paiudosa larvae parasitize Tipulid larvae (Coggins, 1970), which evidently survive the attack (Carter, 1977). The larvae of Borophaga inerassata parasitize Bibionid larvae (Morris, 1922). Species of Phalacrotophora parasitize coccinellid beetle pupae (Van Emden, 1950; Disney, 1979b). Aenigmatias larvae attack ant pupae (Domisthorpe, 1927). Pseudacteon larvae parasitize adult worker ants (Donisthorpe. 1927, Disney 1979b).

It is evident from this brief review of larval habits that we need to be highly critical in our interpretations of rearing records of Phoridae. Species reared from dung, carrion, fungi, or the nests of wasps, bumble bees and other creatures are very likely to be predators or parasitoids rather than saprophages. At the same time species reared from insect 'hosts' found to be dead at the time of collection may indeed be saprophages rather than parasitoids. The polyphagous saprophage species M. sea/aris, M. rujipes and M. giraudii have repeatedly been reported as parasitoids. In no case, however, is the evidence convincing. All three species readily oviposit in the vicinity of moribund insect larvae and pupae and are common pests of insect cultures for this reason. I have found it necessary to keep rearing tubes with phorid larvae and/or suspect hosts enclosed in plastic boxes in order to prevent contamination by these species.

For most species the larval natural history is not known. There is enormous scope for worthwhile investigations by amateur and professional entomologists alike.

Adult natural history is little known. Species of the genus Phora are noted for the swarming habit in the males. These swarms use a marker such as a projecting bough of a tree, a cairn of stones or a person. Some species, notably Borophaga incrassata, can be observed 'drumming' on the leaf on which they are resting. Many species can be observed 'gyrating' and running around on leaves. In autumn adult Phoridae may invade houses in large numbers (Colyer, 1954<1).

Several species visit flowers, particularly Umbelliferae (Disney, 1980a). Diplonevra nitidula has been observed piercing and feeding from a leaf (Disney, 1979b). Fragmentary observations by early workers (e.g. Wesche, 1908) suggest some species may prey upon other insects by piercing and feeding from them. A continental species

6

of Metopina has been observed persuading ants to regurgitate food on which it then feeds (Goetsch, 1940).

Collecting

Any method of collecting insects is likely to procure some species of Phoridae. An these methods are, of course, selective. Thus pitfall traps produce a somewhat poor return for the effort but are an extremely effective way of catching Megaselia /ongicostalis (e.g. Disney et al., 1981). A high return for small effort is given by the use of water traps (bowls part filled with water plus a few drops of detergent). White traps and yellow traps procure the highest diversity of phorids. Malaise traps also produce a relatively high diversity, comparable with that procured in white water-traps. They are poor for some species readily caught in the water-traps but more successful for other species rarely caught in the latter. However a Malaise-trap catch tends to be swamped by large numbers of flies other than Phoridae so that sorting the catch can be tiresome. Sweep-netting is fairly effective but is not only selective in its own way but has a tendency to damage specimens. Netting or pooting scuttle flies off flowers can be very rewarding. Likewise pooting phorids off windows reveals a surprising variety of species, apart from the common house-species Megaselia rujipes. Some species are associated with special situations like the nests of wasps or wood ants. Others are readily procured by the use of emergence traps, for example set over soil (e.g. Disney, Henderson et al., 1981). Some species are most readily procured by rearing from material collected in the field. For example fungus-breeding species are readily procured in this manner. Megase/ia juscinervis has been collected in higher numbers by -collecting shells of its snail host (Vitraea) occupied by puparia than the use of a range of the above collecting methods in the same localities!

Preservation and examination

Previous students ofPhoridae have advocated the micro-pinning of most adults, but have recognised that it may be necessary to slide-mount wingless species and detached pieces (wings, genitalia, etc) (e.g. Borgmeier, 1963). The present author has discovered no merit in micro-pinned specimens. Most of the errors of the past result from the difficulty of discerning detail on a small, usually dark, specimen impaled on a pin. Side­pinning in particular tends to destroy critical characters. Museum collections abound in specimens misidentified because pinning has destroyed or led to the concealment of characters necessary for correct identification. The only character easily observed on most pinned specimens is the detail of the wings. The Phoridae, however, are characteri­sed by a surprising uniformity in the wing venation throughout the family. The differences in wing venation between species tend to be relatively trivial. Furthermore variation within species frequently erodes the taxonomic value of the differences that have been described (see Disney, 1980b).

To add to the sound objections to pinned specimens on grounds of taxonomic acceptability are strong objections from ecologists (e.g. Southwood, 1980). For most ecological purposes preservation of specimens in fluid is greatly to be preferred and in many cases (e.g. with pitfall or water-trap catches) is inescapable. The ecologist is required to handle replicated, quantitative samples of insects in pursuit of his goals. He is frequently confronted with several thousand specimens. To micro-pin such samples would be a labour beyond the resources available to most ecologists. If, therefore, having preserved his specimens in alcohol, he encounters a key based on characters only visible in nicely pinned specimens he responds by leaving his identifications at the family or generic level only. The result has been the loss of much valuable ecological information about particular species as well as the overlooking of a number of British species. The

7

latter omission largely results from restriction of collecting methods to those yielding dry specimens only (see above).

Phoridae should be preserved in fluid. 70'70 alcobol is entirely satisfactory. For long term storage it is probably wise to transfer specimens. when they are thoroughly fixed, to a 1 '70 aqueous solution of propylene phenoxetol. Specimens left in alcohol for many years are liable to discolouration.

Many species can be identified in a watch-glass of fluid placed under a stereo­microscope with a good spotlight. In many cases, however, all or part of the specimen will need to be slide-mounted and examined under a compound microscope. For reference collection purposes slide-mounts are the most satisfactory method of storage. For those not in the habit of slide-mounting insects I would point out that the method recommended below will produce a satisfactory mount in a shorter time than that required to produce an acceptable pin-mount of a specimen.

The mounting medium found to be ideal for Phoridae is the Berlese Fluid advocated for Phlebotomine sand flies by Lewis (1973). The formula is as follows:-. Gum arabic (picked lumps) ......................................... 12g

Chloral hydrate crystals ............................................ 20g Glacial acetic acid .......................... ; ...................... 5m! 50'70 w/w glucose syrup ............................................ 5ml Distilled water ................................................ 30-4OmI Lewis advocates dissolving the constituents at room temperature in the order shown,

filtering the mixture and then evaporating it (at not more than 30°C) until it reaches the required consistency. However, the filtration is not easily achieved satisfactorily. Henshaw (1981) has suggested the impurities in the gum arabic are more easily removed by enclo~ing the latter in a cotton bag and then immersing the latter in the distilled water until the pure gum arabic has passed through the bag into solution. Those not wishing to make up their own medium can purchase it from a dealer (such as GBI Laboratories Ltd, Shepley Industrial Estate, Audenshaw, Manchester, M34 5DW). In view of the number of different formulations of Berlese Fluid that have been evolved it is advisable to specify the formulation given above when placing an order with a dealer.

Berlese Fluid to some extent clears specimens. There is no need to soak Phoridae in a caustic solution before mounting in Berlese Fluid.

For permanent storage slide-mounts in Berlese Fluid need to be dried on a hot plate and then sealed with a ringing medium. A completely trouble-free ringing medium is Glyceel (supplied by BDH Chemicals Ltd (Gurr), Pook, BH12 4NN). Most brands of ladies' nail varnish seem to be almost as satisfactory!

The following conventions ha~been adopted when making a Standard slide-mount of a fully dissected scuttle fly. The dissection is carried out with a pair of fine (watchmaker's) forceps and mounted needles, with the specimen placed in the first drop of Berlese Fluid on the slide beneath a stereo microscope. The specimen is briefly placed on a piece of tissue to remove surplus preserving fluid before it is transferred to this drop of Berlese Fluid. The parts of the dissected fly are arranged as in the figure below:-

+oIL ~ @(t)~ DL lIP' '" ..,; \I) -< '"

.., ~ " 'V ~ '1"' -1 ~ ; 11"'- £ . ...... s: '" ... ~

: ); ~ " OD ~ V . 4' ~. -- ®@ - -.b -u S ~"';."'t7 -.

@ ..u ; -lo. ~

... ~ " ~ 0 r.!..~ , :t IP " .1 __ .. "< r ::; :a A - o ... -

8

The right end of the slide carries the data label (D L) and the left end the identification label OL). The top right mount covers the right wing (rw), the top middle the head (he), the bottom right the right haltere (ha), the bottom middle the front and middle legs (11 & 2), the bottom left the hind legs (13) and finally the top left cover slip covers the rest of the body (thorax and abdomen) (ta) with the left side uppermost. lOmm diameter coverslips are used for all mounts save the haltere, which is covered by a 6mm coverslip (or Cl half a lOmm coverslip). In the case of large species the medium is allowed to dry a little before putting the coverslip over the thorax and abdomen. It has been found easier to avoid air bubbles if a small dab of Berlese Fluid is put on the coverslip's underside before it is lowered over the specimen. In some genera it is necessary to detach the abdomen, at least in one sex, and to mount it dorsal or ventral side upwards. In other cases it is imperative to separate the male hypopygium. All such deviations from the standard mount procedure are indicated at the start of the key to species for the genus in question.

If one needs to re-mount a pinned specimen the most rapid method is as follows: The specimen is soaked in 7011,10 alcohol until it can be carefully slid off its micropin. It is then transferred to Barber's Fluid ( = 33Om195% alcohol, 300ml distilled water, 150ml ethyl acetate, 120ml ether, 10-20 drops of acetic acid) for at least 10 minutes, rinsed in water, and then mounted in the normal way. A simpler, but less rapid method is to soak the specimen in a 1 % aqueous solution of propylene phenoxetol for several days. For some old specimens excessively full of air a soaking in a solution of 2% tri-sodium orthophosphate + 511,10 dimethyl sulphoxide for up to 24 hours, followed by a brief washing in water, has been found to be very satisfactory ( Warning: Do not allow this solution to get on one's skin).

If one needs to examine a feature of a slide-mounted specimen that is on the underside of the mounted part the slide can be turned upside down. It should be supported at either end. If the microscope lacks a slide holder, to act as such supports, a slide placed each side of the stage will serve as well. The specimen can now be viewed in the normal manner, but the reduced clearance will probably prevent use of the highest-magnification objectives.

A compound microscope with a light source whose aperture and intensity can be adjusted as required is to be preferred when examining slide-mounts of Phoridae. Small dark hairs against a dark background may be overlooked if the lighting is incorrectly adjusted. Proper lighting at lower magnification reveals more than poor lighting at higher magnifications!

Morphology

Elsewhere (Disney, 1975) I have commended the views of Tuxen (1969) when he wrote 'the taxonomist should.stick to the terms in common use for his group andno changes should be made for nomenclatural, morphological or semantic reasons'. Such policy means the terms I use are those commended in the introductory volume for these Handbooks on Diptera (Oldroyd, 1970) but out ofIine with more fashionable practice (e.g. McAlpine et at., 1981). Venturi (1966), in justifying his use of terms favoured by some early morphologists, goes too far in regarding the hypotheses of morphologists as being always a scientific bore! However recent work on developmental compartments, recognised from studies on the fate of genetically marked clones of cells labelled during the early stages of development in the embryo of Drosophila (see Palka et al., 1981, and literature cited therein), strongly suggests that much 'established' morphological opinion may be li~le more than flimsy hypothesis. In particular it seems that the identity of the segments (as opposed to compartments) involved in the construction of a particular part of an insect may, in many cases, be secondary to the fact that the particular part persists as a distinctive functional unit. It seems that during evolution

9

the particular segments employed in the construction of the latter may be more opportunistic than has been assumed by many morphologists. Such a view would not be surprising in view of the comparable situation long recognised by students of vertebrates (e.g. fig. 62 in Hardy, 1965).

The HEAD of a scuttle fly is characterised by being dichoptic in both sexes. It typically bears prominent bristles on the frons, which are labelled in fig. 6. The frons may be divided by a MEDIAN FURROW running from the supra-antennal bristles to the anterior ocellus (e.g. figs 124-126). It is, however. frequently reduced to a vestige just above the supra-antennals (e.g. figs 23 and 24) or is absent altogether (e.g. fig 6). In some genera (e.g. Gymnophora) the frontal bristles are reduced or absent. The third antennal segment characteristically embraces and conceals the second segment. The palps usually bear conspicuous bristles (e.g. figs 131-133).

The THORAX has a 'humped' appearance in most species. It bears a variable number of bristles on the dorsum, mainly at the side (figs 9-10) and with usually 2 or 4 bristles on the scutellum. The presence of an inverted L-shaped furrow dividing the mesopleuron is of considerable taxonomic significance. This MESOPLEURAL FURROW is prominent in the middle of the mesopleuron (fig 9). Its upper arm may be crowded dorsally. When supposedly absent (fig. 10) it is possible that elements of this furrow are present but close to the edges of the mesopleuron. Morphological research on the pleural region of the thorax in Phoridae is likely to be rewarding in terms of phylogenetic reconstruction.

The ABDOMEN normally bears tergites on segments 1-6 and may, or may not, bear reduced ones on the remaining segments. However, the first 6 tergites may also be reduced, lost, or modified, in females from the posterior end forwards (e.g. in Diplonevra and Metopina). Typically sternites are lacking on the first six segments, but a distinct stemite 6 is present in Pseudacteon females and an often ill-defined plate is present on the venter of Metopina males (figs 79-80). The stemite of segment 7 is frequently present in females and in Triphleba is of considerable taxonomic importance (figs. 166-176).

The male HYPOPYGIUM may be very complex. The principal visible parts are labelled in fig. 7. The penis complex may be fully withdrawn. When fully extruded it may display an elaborate structure of considerable complexity (e.g. fig. 155). The anal tube may be much shortened or greatly elongated. The epandrium frequently bears conspicuous processes (e.g. in Triphleba, figs 143-144, 150-154). In females the terminal segments may be developed into a relatively simple ovipositor (e.g. figs 99-100) or modified into a sharp homy structure, associated with parasitic habits,(e.g. in Pseudacteon) .

The WINGS are highly characteristic in the simplification of their venation. The latter is surprisingly uniform throughout the family and is labelled in fig. 8. The homologies of the thick veins are still disputed and so are labelled 1-3, without reference to which radial veins they might represent. The wing length is measured from the basal bristle to the wing tip, but along a line parallel to the costa from the wing tip. The costal index (costa length, from basal bristle, divided by wing length) has not been used in the present keys. Likewise the costal ratios (costal section 1 divided by costal section 3, section 2 divided by section 3) do not require to be measured in the present keys. Wings are absent in the females of some genera (e.g. figs 15 and 29).

The LEGS frequently carry isolated bristles on the tibiae apart from the apical spurs. Many have one (or sometimes more) dorsal hair-palisades (e.g. fig. 12) on the middle and hind tibiae. In some there are overlapping transverse combs of stout hairs instead (e.g. fig. 75). The posterior face of the base of the hind femur of males frequently has features of considerable taxonomic value (e.g. in Diplonevra, Dohmiphora, Gymnoptera and Metopina).

lO

Notes on the keys

The keys are based upon the examination of slide"mounted specimens and I have assumed that if a detail cannot be seen dearly under low magnifications it will be examined as a slide-mount with a compound microscope.

The keys represent a radical departure from previously published keys. On top of this I have made much greater use of illustrations to clarify the details at issue. Throughout the figures I have used a scale line of 0.1 mm, so that the degree of magnification is immediately apparent.

In some cases (e.g.linthelkeyltoIPhora !species) the key has been designed in such a way that an answer may be reached by more than one route. This is to allow for variation in otherwise useful characters (like the number of bristles on the tibia). In other cases I have keyed a species twice if an otherwise useful character varies in a particular species.

In some cases (e.g. Aenigmatias and Phora) the females are not keyed to species as present knowledge does not allow this.

In preparing these keys, I have added a dozen species not previously recorded for Britain. There is, therefore, a high probability of other species turning up. I will gladly examine any specimens that do not appear to be included in the present handbook. Otherwise the following works should be consulted:- Schmitz (1938b-1958, continued as Schmitz & Beyer, 1965-1974, Schrnitz& Delage, 1974-1981) and Lundbeck (1922). In addition the European Pseudacteon are covered by Schmitz (1938a), Metopina by Disney (1979j), and Phalacrotophora by Delage & Lauraire (1974) and Disney (1979k).

Information on distribution is based primarily on my own records. This is supple­mented from published information, museum collections and Colyer's unpublished notebooks. However these secondary sources of information have been used with extreme caution, particularly in genera (e.g. Phora and Metopina) where remounting on slides of museum specimens on pins has revealed a high frequency of misidentifica­tions and/or revisionary work has revealed unsuspected species (e.g. Gymnophora). Likewise the data on adult phenology (expressed by the use of Roman Numerals for months for which there are records) are based primarily on my own records. The most useful reference collection of British Phoridae is my own collection of slide-mounted specimens!

In most genera the sexes can only be separated on the basis of the abdominal terminalia (see under 'Morphology' above).

Acknowledgements

I am much indebted to the late Mr. C.N. Colyer whose translation from the German of Schmitz's keys to the Palaearctic Phoridae saved much time with a dictionary. In addition his notebooks (deposited in the British Museum - Natural History) have been a valuable source of information.

K.G.V. Smith, who originally intended being co-author (before his duties obliged him to drop out), has greatly encouraged me. He and B.H. Cogan have freely allowed access to the collections (of the British Museum - Natural History) in their care and most generously allowed me to re-mount critical specimens on slides. Likewise Dr. H. Ulrich (Zooiogisches Forschungsinstitut and Museum Alexander Koenig, Bonn) has been more than generous with the loan of specimens from the Schmitz Collection. Dr. M.C. Birch (Hope Entomological Collections, Oxford University), Dr. M.R. Shaw (Royal Scottish Museum) and J.P. O'Connor (National Museum of Ireland) have also loaned valuable specimens in their care.

Mrs B.G. Leonard (Royal Entomological Society Library) as well as Pamela Gilbert and Ann Lum (Entomology Library, British Museum - Natural History) have been

11

tireless in searching out obscure pUblications for me. I must acknowledge the numerous people who have sent me collections of Phoridae

nicely preserved in alcohoL In particular valuable specimens were received from E.C.M. d'Assis Fonseca, Dr. M. Boness, P.J. Chandler, Dr. J.C Coulson, Dr. L. Davies, D. Henshaw, D. Howse, P. Holden, Dr. A.G. Irwin, Dr. J. Ismay. Dr. LF.G. Mclean, J.M. Nelson, P. Skidmore, D.A. Smith, Prof. T.R.E. Southwood, A.E. Stubbs, Dr. M.CD. Speight, R. Szadziewski, D.M. Unwin and P. Withers.

I am much indebted to the Royal Society and to the Shell International Petroleum Co. Ltd for grants without which the preparation of this handbook would have taken longer.

Finally, the Trustees of the British Museum (Natural History) are thanked for the use of the drawing of the Megaselia sealaris female used on the front cover. This drawing was executed by Arthur Smith.

Check list

The following list corrects and updates that in Kloet & Hincks (1976). The species of Megaselia are omitted as they will be covered in a separate Handbook. The species in the genus Plastophora have been returned to the genus Megase/ia (Disney, 1978). The species of Citrago are now placed in Triphleba (Disney, 1982g). The detailed documentation of the other additions and corrections is to be found in Smith (1977) and Disney (1979 a and c, 1980 doe, 1981 b-c. e-f, 1982 a-b, d-i).

The total of 93 species in 21 genera are listed below. The genera are arranged alphabetically and are not placed in subfamilies as the definitions of the latter are currently in question.

PHORIDAE

AENIGMATIAS Meinert, 1890 PLATYPHORA Verrall, 1877

brevifrons Sehmitz, 1955 lubbocki (Donisthorpe, 1913) nee

(Verrall, 1877) dorni (Donisthorpe, 1914) nee,

(Enderlein, 1908) jranzi Sehmitz, 1950 lubbocki (Verrall, 1877)

blattoides Meinert, 1890 v.highlandicus Sehmitz, 1914

ANEVRINA Lioy, 1864 ANEURINA, emend. auett., nee

Sehmitz, 1941 cUn.>inen.>is (Beeker, 1901) thoracica (Meigen, 1804)

dimidiata (Meigen, 1830) unispinosa (Zetterstedt, 1860)

jennica (Beeker, 1901) urbana (Meigen, 1830)

quadrata (Gimmerthal, 1842) trochanterata (Zetterstedt, 1855) caliginosa (Malloeh, 1912) nee

(Meigen, 1830)

BECKERINA Malloch, 1910 umbrimargo (Beeker, 1901)

12

BOROPHAGA Enderlein, 1924 agi/is (Meigen, 1830) carinifrons (Zetterstedt, 1848) jemorata (Meigen, 1830)

j/avimana (Meigen, 1830) incrassata (Meigen, 1830) irregularis (Wood, 1912) subsultans (Linnaeus, 1766)

jemorata (Brues, 1904) nee (Meigen, 1830)

j/avimana (MaIloeh, 1912) partim nee (Meigen, 1830)

o'kellyi Sehmitz, 1937

CHAETOPLEUROPHORA Sehmitz, 1922

bohemanni (Beeker, 1901) erythronota (Strobl, 1892)

v.nigrodorsata (Strobl, 1910) spinosior Sehmitz, 1938 spinosissima (Strobl, 1892)

CHONOCEPHALUS Wandolleck, 1898

heymonsi Stobbe, 1913 jamaicensis Brues, 1915 punctifascia Borgmeier, 1935 vadoniPaulian, 1958

CONICERA Meigen, 1830 dauci (Meigen, 1830)

albipennis (Meigen, 1830) pro parte

atra Meigen, 1830 nickerli (Kowarz, 1894)

jloricola Schmitz, 1938 similis Schmitz. 1920 partim nee

(Haliday, 1833) similis var. Julvipalpis Sehmitz,

1920 minuscula Schmitz, 1953

schnittmanni Schmitz 1926 similis (H!!1iday, 1833)

pauxilla Schmitz, 1920 tarsalis Sehmitz, 1920 tibialis Sehmitz, 1925

albipennis (Meigen, 1830) pro parte

similis Schmitz, 1924 nee (Haliday, 1833)

Jallens Sehmitz, 1948

DIPLONEVRA Lioy. 1864 DlPLONEURA (emend.) Schmitz,

1929 abbreviata (von Roser, 1840)

sordidipennis (Dufour, 1841) concinna (Meigen, 1830)

crassicornis (Zetterstedt, 1848) nee (Meigen, 1830)

amplicornis (Schmitz, 1926) jlorea (Fabricius, 1794)

abdominalis (Fallen, 1823) palpina (Zetterstedt, 1848) Julviventris (Boheman, 1852) jlexuosa (Egger, 1862) sororcula (Van der Wulp, 1871) versicolor (Sehmitz, 1920)

Junebris (Meigen, 1830) atra (Maequart, 1835) nee

(Meigen, 1804) pseudoconcinna (Strobl, 1892) cimbicis (Aldrieh, 1892) luctuosa (Becker, 1901) concinna (Malloeh, 1912) nee

(Meigen, 1830) rostralis (Sehmitz, 1918) nitidula (Sehmitz, 1929) nee

(Meigen, 1830) glabra Sehmitz, 1927

parcepilosa Sehmitz, 1927 nitidula (Meigen, 1830)

Juctuosa (Meigen, 1838) gymnophorina (Z~tterstedt, 1848) distincta (Egger, 1862) concinna (Beeker, 1901) nee

(Meigen, 1830) nitidifrons (Brues, 1904)

pilosella Sehmitz, 1927

DOHRNlPHORA Dahl, 1898 corn uta (Bigot, 1857)

navigans (Frauenfeld, 1867) cleghomi (Bigot, 1890) venusta (Coquillett, 1895) chlorogastra (Beeker, 1901) mordax (Brues, 1911) bequaerti Sehmitz, 1915 jlaviventris (Silva Figueroa, 1916) divaricata v. basalis Santos Abreu,

1921 divaricata v. obscura Santos Abreu,

1921 Julva Santos Abreu, 1921 opposita Borgmeier, 1925 crockeri Van Duzee, 1933 willowsi Van Duzee, 1933

GYMNOPHORA Macquart, 1835 arcuata (Meigen, 1830)

rufipes (Fallen, 1823) partim nee (Meigen, 1804)

debilis (Haliday, 1833) healeyae Disney, 1980 integralis Sehmitz, 1920 quartomolIis Sehmitz, 1920

GYMNOPTERA Lioy, 1864 longicostalis Sehmitz, 1933

vitripennis (Wood, 1906) nee (Meigen, 1830)

vitripennis (Meigen, 1830) genitalis Sehmitz, 1927

HYPOCERA Lioy, 1864 mordellaria (Fallen, 1823)

jlavipalpis (Maequart, 1835) subsultans Sehmitz ante 1940 nee

(Linnaeus, 1766)

MEGASELlA Rondani, 1856 Not included in this handbook

METOPINA Macquart, 1835 braueri (Strobl, 1880)

cuneata Sehmitz, 1924 crassinervis Sehmitz, 1920 galeata (Haliday, 1833)

inaequalis Sehmitz, 1927 heselhausi Sehmitz, 1914 oligoneura (Mik, 1867)

galeata Lundbeek, 1922 partim nee (Haliday, 1833)

nevadae Sehmitz, 1957 perpusiIla (Six, 1878)

gaieata Sehmitz, 1936 nee (Haliday, 1833)

13

rhenana Beyer and Schmitz, 1957 pileata Sehmitz, 1936 ulriehi Disney, 1979

PHALACROTOPHORA Enderlein, 1912

berolinensis Sehmitz, 1920 jasciata (Fallen, 1823)

atrieapilla (Curtis, 1833) nigrocineta (de Meijere, 1907)

PHORA Latreille, 1796 artijrons Sehmitz, 1920

stietlea (Zetterstedt, 1848) partim nee (Meigen, 1830)

atra (Meigen, 1804) aterrima (Fabricius. 1794) preoee.

(Villers, 1789) bullata Sehmitz, 1927 dubia (Zetterstedt, 1848)

stietiea (Zetterstedt, 1838) nee (Meigen, 1830)

sehineri (Seeker, 1901) edentata Sehmitz, 1920 hamata Sehmitz, 1927 holoserieea Schmitz, 1920

aterrima (Zetterstedt, 1848) partim nee (Fabricius, 1794)

velutina de Meijere, 1919 nee (Meigen, 1830)

obscura (Zetterstedt, 1848) aterrima (Zetterstedt, 1848) partim

nee (Fabricius, 1794) heteroeerea Sehmitz, 1920 hamata Sehmitz, 1947 nee

Sehmitz, 1927 praepandens Sehmitz, 1927 speighti Disney, 1982 stietiea Meigen, 1830

velutina (Beeker, 1901) nee (Meigen, 1830)

montana (Brues, 1904) tineta Sehmitz, 1920

PLECT ANOCNEMA Sehmitz, 1926 nudipes (Beeker, 1901)

PSEUDACTEON Coquillett, 1907 brevieauda Sehmitz, 1925 jormiearum (Verrall, 1877)

minor (Strobl, 1880) vitripennis v. albohalterata

(Strobl, 1892)

PULICIPHORA Dahl, 1897 borinquenensis Wheeler, 1906

oecidentalis Barber, 1905

14

ajrieana Brues, 1907 spinieollis Schmitz, 1915

termitophila Seevers, 1941 wymani Bohart, 1947 nuptamjerens Paterson, 1953

SPINIPHORA Malloeh, 1909 bergenstammi (Mik, 1864)

maculata v. immaeu/ata (StrobL 1880)

sphingicides (Strobl. J 892) nee sphigicides (Bouehe, 1834)

domestiea (Wood, 1906) pressata (Beeker, 1919) intrieata (Sehmitz, 1935)

dorsalis (Beeker, 1901) thoraciea v. immaeulata

(Strobl. 1894) exeisa (Beeker, 1901)

bergenstammi (Seeker, 1901) partim nee (Mik, 1864)

eomstoeki (Aldrieh, 1904) maculata (Meigen, 1830)

vulgaris v. "forte spec. dist." (Fallen, 1823)

jlavieornis (Maequart, 1835) helicivora (Dufour, 1841) notata (Zetterstedt, 1848)

TRIPHLEBA Rondani 1856 CITRAGO Sehmitz, 1924

antrieola (Sehmitz, 1918) aptina (Bezzi, 1911) nee

(Sehiner, 1853) bartholomei (Sehmitz, 1921) jantinii Sehmitz, 1933 eoncolor Sehmitz, 1940 nom. nudo

autumnalis (Beeker, 1901) hiemalis (Dahl, 1896) jorcipata (Strobl, 1910)

citreijormis (Beeker, 1901) eollini Sehmitz, 1943 crassinervis (Strobt, 1910)

enervata (Sehmitz, 1920) distinguenda (Strobl, 1892)

unispinosa (Seeker, 1901) uniealearata (Seeker, 1901) albispina Boheman (Seeker. 1901)

nom. nud exeisa (Lundbeck, 1921)

oetobris (Schmitz, 1921) j1exipalpis Sehmitz, 1927 gracilis (Wood, 1907) hyalinata (Meigen, 1830)

perennis (Meigen, 1838) hyemalis Rondani 1856 opaca (Beeker, 1901) nee

Meigen, 1830) perennijormis Sehmitz, 1934 mohrae Sehmitz, 1938

intempesta (Schmitz, 1918)

intermedia (Malloeh, 1908) lugubris (Meigen, 1830)

fratercula (Brues, 1904) sublugubris (Wood, 1906) suspecta (Malloeh, 1912) connexa (Wood, 1914)

luteifemorata (Wood, 1906) similis (Lundbeck, 1921)

minuta (Fabricius, 1787) ru/icornis (Zetterstedt, 1848) nee

(Meigen, 1830) pubericornis (Malloeh, .1908)

nudipaipis (Beeker, 1901) opaca (Meigen, 1830)

nigricomis (Egger, 1862) lehmanni (Enderiein, 1927)

papillata (Wingate, 1906) lugubris (Wingate, 1906) nee

(Meigen, 1830)

opaca (Sehiner, 1864) partim nee (Meigen, 1830)

smithi Disney, 1982 disparinervis (Schmitz, 1949)

partim nee (Sehmitz, 1947) collini (Schmitz, 1955) preoec.

collini Schmitz, 1943 subcompleta Schmitz, J 927 trinervis (Beeker, 1901)

urbana (Meigen, 1830) pro parte opaca (Sehiner, 1864), nee

(Meigen, 1830) gilsoni (Sehmitz, 1915)

vitrea (Wood, 1906)

WOODIPHORA Sehmitz, 1926 retroversa (Wood, 1908)

Key to genera

1 Wingless and without halteres (female only) ..................................... 2 With wings and halteres (both sexes) ........................................... 4

2 Fifth abdominal tergite divided into two parts. The anterior part is a semi-circu1ar flap attached only along its straight, anterior edge (fig. 128). Ocelli present.PULICIPHORA (p.31)

Fifth abdominal tergite undivided. Ocelli absent. ................................ 3 3 The single thoracic notum broader and longer than head and with convex anterior margin

(fig. 15). Frons with small hairs only. Head short but broad (fig. 15) .............. . ............................................... . AENIGMATIAS (p. 17)

The single thoracic notum about as broad but clearly shorter than head and with both anterior and posterior margins somewhat concave (fig. 29). Frons with some bristle like hairs at sides in front of eyes, as well as small hairs on rest of frons. Head only a little broader than long (fig. 29) ................................. CHONOCEPHALUS (p. 20)

4 Hind tibia with transverse combs, of stoutish hairs, arranged one below the other in an overlapping series along dorsal face (fig. 75). Mesopleuron with hairs and a single long bristle near posterior margiu (similar to fig. 9) ........................ ; ........ 5

Hind tibia either simply haired or with longitudinal palisades of stoutish hairs, apart from any isolated bristles. Mesopleuron frequently without hairs or bristles ............. 6

5 Vein 3 forked at tip (Le. vein 2 is present) and with a single hair at its base on the dorsal face ........................................... CHAETOPLEUROPHORA (p. 20)

Vein 3 unforked and with row of 3-10 hairs extending from its base along the dorsal face ........................................................ HYPOCERA (p. 25)

6 Vein 3 with a series ofat least 5 hairs (which may be rather rme) along its length on the dorsal face (e.g. figs 21 and 22) .................................................... 7

Vein 3 with 0-3 hairs (usuallyOor 1) restricted to basal quarter, or (rarely) with a bristle-like hair at base and 1-4 finer hairs before fork of vein 3 ........................... 10

7 Tibiae with apical bristles only. apart from hairs and hair palisades ................ . . .... . .... . . .. . .............. . .. . . ... . .. . . ... . . . AENlGMATIAS (p. 17)

Most tibiae with one or more isolated bristles in upper two-thirds, as well as apical bristles .......................................................................... 8

8 Hind tibia with a dorsal, longitudinal palisade-like row of stoutish hairs (similar to fig. 12) in addition to isolated bristles and scattered hairs. Vein 3 frequently unforked ..... .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . BOROPHAGA (p. 19) Hind tibia simply haired on dorsal face. Vein 3 always forked (i.e. vein 2 always clearly present)

......................................................................... 9 9 With less than 10 hairs on dorsal face of vein 3 ................................. 10

With more than 10 hairs on dorsal face of vein 3 ................. ANEVRlNA (p. 18)

15

10 Vein 6 (3rd thin vein) with a sudden bend near middle, opposite (and opposed to) basal curve of vein 5 (figs. 76-78) ...................................... METQPINA (p. 25)

Vein 6 without such a bend .................................................. 11 11 Hind tibia with one or more dorsal longitudinal palisade-like rows of stoutish hairs ('p' in

fig. 12) ............ '" ................................................... 12 Hind tibia, apart from any isolated bristles, simply haired on dorsal face (similar to fig. 30).

Seen in profile these hairs are not as densely packed (side by side) as in a palisade row · ................................ " ................................. 19

12 Most tibiae with one or more isolated bristles in upper two-thirds as well as apical bristles. If these bristles are small and inconspicuous check that the mesopleural furrow is apparently absent (fig. 10) ........................................................... 13

Tibiae without isolated bristles, apart from apical bristles, but there may be a row of bristJe­like hairs either side (posterodorsal and antero-dorsal) of the dorsal hair palisades on the middle and hind legs. If in doubt the mesopleural furrow should be examined. It is centrally placed, typically seen as a line running posteriorly from the most posterior point of propleuron and then turning ventrally half way across the mesopleuron (fig. 9) .... 14

13 Mesopleuron bare. Hind tibia with 2, or (rarely) 3, dorsal hair palisades ............. . · ................................................ DIPLONEVRA (p. 22)

Mesopleuron with some hairs on upper part. Hind tibia with a single dorsal hair palisade .................................................... DOHRNIPHORA (p. 23)

14 Vein 3 unforked (i.e. vein 2 is missing) .. " ................... , ................. 15 Vein 3 forked at tip ......................................................... 16

15 Metatarsus of front leg slightly shorter than fifth segment. 9 with distinct sternite on segment 6 of abdomen and a down-curved pointed ovipositor behind ..................... .

· ............................................... PSEUDACTEON (p. 30) Metatarsus of front leg clearly longer than fifth segment. Abdominal venter devoid of sternites

on segments 1-6 in both sexes. (aberrant specimens with vein 2 missing) ........... . · ............................ MEGASEUA (not included in this handbook)

16 Hind tibia with a series of somewhat irregular hair palisades (fig. 26) ................ . ........................................ ... . PLECTANOCNEMA (p. 30)

Hind tibia with a single, more-or-less dorsal, hair palisade (similar to fig. 12) ........ 17 17 Hind tibia with an antero-dorsal as well as a postero-dorsal row of bristle-like hairs (with

hair palisade running between the two rows) .................................. 18 Hind tibia with no antero-dorsal row of bristle-like hairs. The postero-dorsal row always

present, but the hairs may be somewhat fine ................................... . ........... ................. . MEGASELIA (not included in this handbook)

18 Palp bristles only about size of upper occipitals. 9 , with tergites present only on abdominal segments 1-4 (the fifth segment dorsally with a complex gland opening). Tergite 3 with a pair of small, oval to round papillae - one to each side towards front margin. er, with proctiger ending in finely-feathered bristles which are clearly more robust than hairs on cerci (figs. 97 and 98) ........................... PHALACROTOPHORA (p. 28)

Palp bristles strongly developed, the longest being clearly more robust than upper occipitals. 9 , with tergites present on abdominal segments 1-6 (in M. melanocephala a gland opens between tergites 5 and 6) and without papillae on tergite 3. er, with proctiger ending in hairs little, if any, stronger than those on cerci (i.e. weaker than those in fig. 7) ..... .

· ............................ MEGASELIA (not included in this handbook) 19 Most tibiae with one or more isolated bristles in upper two-thirds as well as apical bristles

(e.g.figs 40-43). If these bristles are small and inconspicuous check that the mesopleural furrow is apparently absent (fig. to) ......•..•............................... 20

Tibiae without isolated bristles, apart from apical bristles (e.g. fig. 30). Mesopleural furrow centrally placed (being seen as a line running posteriorly from the most posterior point of propleuron and thelHurning ventrally half way across the mesopleuron - fig. 9) ..

................................................................... . 26 20 Vein 3 forked at tip (e.g. figs. 145-149) ....................... '" .............. 21

Vein 3 unforked (Le. vein 2 missing) (e.g. figs. 25 and 127) ....................... 23 21 Mesopleuron with hairs and a single bristle near posterior margin (similar to fig. 9) ... .

· ..................................... CHAETOPLEUROPHORA (p. 20) Mesopleuron bare .......................................................... 22

22 Fore tibia with a bristle in upper two-thirds which is longer than maximum breadth of tibia,

16

and with a row of short, robust, near-dorsal spines below this bristle (fig. 11). First two tarsal segments (of foreleg) with a clearly differentiated, short, dorsal apical spur ('5' in fig. 11). Mid-tibia with anterior bristle in distal half which is clearly longer than width of tibia at point of insertion .............................. , .SPINIPHORA (p. 31)

Fore tibia with bristle in upper two-thirds usually shorter than maximum breadth of tibia or absent, and usually without a clearly differentiated row of short, robust, antero-dorsal spines below this bristle. First two tarsal segments (of fore leg) without differentiated, short, dorsal apical spurs. Mid tibia either without anterior bristle in distal half or if a pre-apical bristle is present its length is usually less than width of tibia at point of insertion .....

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . TRIPHLEBA (p. 32) 23 Mid tibia with at least 2 true-dorsal bristles. Frons with a complete median furrow (figs.

124-.126). (Very black species) ..... , ............................. PHORA (p. 28) Mid tibia at most with 1 true-dorsal bristle. Frons without median furrow (e.g. fig. 6) (There

may be a slight indication of a furrow restricted to the region of the supra-antenna! bristles) ........................................................................ 24

24 Hind tibia with at least 2 bristles in upper two-thirds. Arista inserted apically ........ 25 Hind tibia at most with 1 bristle in upper two-thirds. Arista inserted dorsally in a pre-apical

position ................................................. TRIPHLEBA (p. 32) 25 Vein Se ends before reaching vein I. Hind tibia with at least 2 true-dorsal bristles and 1 antero­

dorsal near the upper one (figs. 40-43). et, third antennal segment drawn out into elongated neck ..................................................... CONICERA (p. 21)

Vein Sc runs into (and coalesces with) vein 1. Apart from apical bristles hind tibia with 2 antero-dorsal bristles only, one situated near end of upper third and one near middle. et, Third antennal segment only slightly pointed distally ....... GYMNOPTERA (p. 25)

26 Vein 3 forked at tip (figs 20,69 and 179) ....................................... 27 Vein 3 unforked (Le. vein 2 missing) (figs 25 and 127) ............... , ............ 29

27 Frons without bristles between anterior ocellus and base of antennae, but with numerous short hairs. Mesopleuron with a patch of fme hairs near dorsal edge (fig. 65) (wings brownish or strongly greyish) ................................... GYMNOPHORA (p. 24)

Frons with conspicuous bristles, between anterior ocelIus and base of antennae, as well as short hairs (similar to fig. 6). Mesopleuron bare ..................... ; ......... 28

28 Costal cilia clearly longer than vein 2 (proximal branch of fork of vein 3). Costa thickens somewhat distally (fig. 20). Wing membrane conspicuously darkenedBECKERlNA (p. 19)

Costal cilia shorter than vein 2. Costa not thickened (fig. 179). Wing membrane only slightly greyish ............................................... WOODIPHORA (p. 37)

29 Base of vein 4 (1st thin vein) obliterated. Vein 3 ending in a small, clear vesicle just beyond the point where it meets the costa. The angle between the costa and end of vein 3 is small (fig. 25). Wing membrane clearly greyish or greyish brown ...................... .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . CHONOCEPHALUS (p. 20) , Base of vein 4 complete. Vein 3 not ending in such a vesicle and meeting costa at a wider

angle (fig.127). Wing membrane only slightly greyish ....... PULICIPHORA (p. 31)

Genus Aenigmatias Meinert (Figs. 13-17)

This genus includes at least a dozen species, of which 7 are known from Europe. The other species occur in the Nearctic Region.

The larvae are known to parasitize the pupae of ants, particularly wood ants (Formica spp.). The genus is poorly known in Britain and early observations probably confused the different species now known to occur. There are probably other species awaiting addition to the British List. Only the males are keyed below as the females of two species are not yet known. The author would welcome specimens (in alcohol) of this genus.

In order to avoid overlooking previously unrecorded species it is advisable to tease out the hypopygial capsule and mount it as in fig. 16. (It should be noted that the wing venation characters traditionally used to distinguish A. lubbocki from related species are unreliable, due to variation that has not been allowed for).

17

Key to species Males

Basal half of hind femora more or less yenow, likewise the middle femora. (Wing as fig. 14. Hypopygium as figs. 16 and 17.) ............................... Iubbocki (Verrall) Hants., Hereford, Scotland; Inverness. vii-ix.

All legs a uniform dark brown to almost black in colour .......................... 2 2 Wing membrane uniformly greyish. Veins 4-7 all pale and difficult to discern at low

magnifications (x to or less) ................................. brevifrons Schmitz Surrey. vii.

Wing membrane grey with some areas brownish tinged, notably in costal cen, against the posterior margin of vein 3 and either side of veins 4-6, which are brown and easily discerned at low magnifications (fig. 13) ................................... frallzi Schmitz Cheshire, Cumbria., Scotland: Inverness vii. viii.

Genus Anevrina Lioy (Figs 18-19)

This genus includes more than a dozen species, of which 5 are Palaearctic, the rest being from the Nearctic and Oriental Regions.

The larvae are associated with carrion of small vertebrates or the soil, particularly mole nests. Detailed studies of larval natural history are lacking.

Keys to species Males

Anterior face of hind femur with an oval patch of minute hairs in basal half (fig. 18). Apart from tip of hind femur and hind tibia the legs are yellowish. (Hind tibia with a pair of bristles at end of basal quarter, a bristle near mid-point, and a pre-apical in distal quarter) · ......................................................... tboracicll(Meigen) Widespread in Great Britain. Ireland: Offaly. v-x.

Anterior face of hind femur without such a patch of minute hairs. Legs brownish to dark brown ................................................................... 2

2 Knob of haltere dark brown to black. Posterior face of base of hind femur with short black spines, some of which are blunt ended (fig. 19) ................ curvinervis (Becker) Widespread in England. S. W. Scotland. iv-v.

Knob of haltere yellow to pale brownish yellow. Posterior face of base of hind femur with fine hairs or short, fmely tapered, pale almost spine-like hairs .................... 3

3 Apart from apical spurs hind tibia with no dorsal bristles and only 2 anterior bristles (1 in basal half and I pre-apical). Anal tube terminal in position and projecting posteriorly. · .....•............................................... unispinosa (Zetterstedt) Widespread in England. Scotland: Dunbarton, Inverness, Renfrew. Ireland: Offaly. v-x.

Hind tibia with 4-8 bristles (apart from apical spurs) of which at least 2 are situated dorsally. Anal tube situated dorsally, in sub-terminal position, and not projecting posteriorly · ........................................................... urbana (Meigen) Widespread in England. Scotland: Dunbarton. iii-vii.

Females (For distribution and phenology see males)

W'mg with distal third or more darkened (brownish grey), contrasting with rest of wing which ·is pale grey to almost clear. Legs yellow to brownish yellow. Abdominal tergite S reduced and with a gland complex that includes an evertible dark-walled sac each side · ......................... ~ ............................... thoraeica (Meigen)

Wing without a contrasting darker distal third. Legs largely brown with only the front legs yellowish brown in some cases. Abdominal segment S without gland complex ...... 2

2 Hin~ tibia with <k8 bristles (apart from apical spurs) of which at least 2 are situated dorsally.

18

Abdominal tergite 5 represented by hairs only .................... urbana (Meigen) Hind tibia with 2-3 bristles (apart from apical spurs) of which one at most is situated dorsally.

Abdominal tergite 5 normally developed ...................................... 3 3 Knob of haltere dark brown to black. Hind tibia with 2 anterior bristles and a single dorsal

bristle below the upper anterior .............................. curvinervis (Becker) Knob of haltere yellow to dusky yellow. Hind tibia with 2 anterior bristles only (1 in basal

half and 1 pre-apical) ................................... omspinosa (Zetterstedt)

Genus Beckerina Ma110ch (fig. 20)

This genus includes nearly 20 species, most of which are Nearctic or Neotropical. One is known from Burma, one from New Zealand and one from Europe.

The larval natural history is unknown.

Wing as in fig. 20. Haltere dark brown to black. All legs dark. Hairs beneath base of hind femur shorter than those on anterior face. <Y, Right side of epandrium drawn out into sub-triangular process extending beyond end of anal tube and bearing hairs which posteriorly tend to be bent over at tip. 9. with abdominal tergite 6 ill-defined and less pigmented than preceeding tergites ......................... umbrimargo (Becker) Widespread in lowlands of England and Wales. Scotland: Dunbarton. Ireland: Kerry. iv­viii.

Genus Borophaga Enderlein (Figs. 21~24)

This genus includes more than two dozen species, of which 10 are Palaearctic, 8 from Burma and the rest from the Nearctic, Neotropical and Afrotropical Regions.

The larvae of B. incrassata parasitize the larvae of Bibio marci (Bibionidae).

Key to species Males and females

Tip of vein 3, from vicinity of origin of vein 4, expands to form a distinctly swollen distal portion (figs. 21-22) .......................................•................ 2

Tip of vein 3 either parallel-sided or only expanding very gradually and even then not forming a distinctly swollen distal portion ........................................... .4

2 Mid-tibia with pre-apical anterior bristle weakly developed so that it is clearly shorter than lower apical spur and is scarcely longer than antero-dorsal spines. Vein 3 is forked, although the anterior branch (vein 2) is very fine and pale (fig. 21.) ........ irregularis (Wood) Devon, Gloucs., Hereford, N. Yorks. Scotland: Sutherland. vii-x.

Mid-tibia with pre-apical anterior bristle clearly longer than lower apical spur. Vein 3 unforked (i.e. vein 2 missing) ...............•..............................•......... 3

3 Hind-tibia with dorsal apical spur equal to or a little longer than the anterior prc-apical bristle (these two bristles are separated by the deflected extremity of the hair palisade).Some of the hairs of mesopleuron situated less than length of a hair from posterior margin ...............................................•.......... femorata(Meigen) Cheshire, E. Anglia, S. of England. Scotland: Dunbafton iii-v, vii-ix.

Hind tibia with dorsal apical spur greatly reduced so that it is scarcely longer than hairs of hair palisade. Hairs of mesopleuron not extending so far back, so that tips of the most posteriorly situated hairs do not quite reach posterior margin. (Tip of vein 3 as fig. 22) ....................................................... subsultans (Linnaeus) Cumbria, Hereford, Hants., N. Yorks. Widespread in Scotland. Ireland: Kerry, Offaly. iii-x.

19

4 Ocellar triangle expanded into an elevated protruberance, and anterior ocenus transversely elongated so that it is at least 3 x broader than high (figs. 23-24). Hind tibia with a strong anterior bristle in upper half. Vein 3 unforked ................................. 5

Ocellar triangle not developed as an elevated protuberance, and anterior ocellus less than twice as broad as high. Hind tibia without bristles in upper two-thirds. Vein 3 usually forked, but the anterior branch (vein 2) tends to be obscure, often incomplete and sometimes absent ..........•.................................................. agilis (Meigen) Berkshire. viii.

5 Elevated protuberance bearing ocelli broader (fig. 23). Front tibia with at least 2 postero-dorsal bristles. In both sexes the cerci are broadly ovoid and covered in densely-crowded fine hairs ......... '" ....................... " ............ mcrassata (Meigen) Widespread in lowlands of England. Ireland: Wicklow. vi-ix.

Elevated protuberance bearing ocelli narrower (fig. 24). Front tibia with only 1 postero-dorsal bristle. In both sexes the cerci are elongated and bear a few robust hairs, some of which greatly exceed the length of the cerci. ..................... carinifrons (Zetterstedt) Widespread in England and Scotland. vi-ix.

Genus Chaetopleurophora Schmitz This genus includes more than 20 species, of which 6 are known from Europe, with rest

being from the Nearctic, Neotropical and Oriental Regions. The larvae have been recorded in dead snails.

Key to Species Males and females

Hind tibia with only 2 bristles, both antero-dorsal, apart from conspicuous apical spurs. Dorsal face of hind tibia with transverse combs (as in Hypocera-fig. 75) ................. 2

Hind tibia with 7 or more bristles, apart from conspicuous apical spurs. Dorsal face of bind tibia without transverse combs ...................•........................... 3

2 Apart from apex of coxae, trochanters and tips of femora the legs are dark brown to black. Mid-tibia with 2 large apical spurs. Thorax always dark ......... bohemanni (Becker) Bucks. (one record only - obtained by rearing)

front and middle legs more or less yellowish. Hind legs with distal half of femora and tibiae dark brown to black. Mid-tibia with only I large apical spur. Thoracic dorsum varies from dark brown to pale orange .................................. erythronota (Strobl) Widespread in England and Wales. S. W. Scotland. Ireland: Clare, Offaly. vi-ix.

3 Mid-tibia with 4 bristles, apart from apical spurs. Hind tibia with only 1 pre-apical anterior bristle (situated in dorsal half) ................................. spinosior Sclunitz Ireland: Clare. Offaly. v-vi.

Mid-tibia with 5-7 bristles, apart from apical spurs. Hind tibia with 2 pre-apical anterior bristles (the lower being shorter than the one above it) ................ spinosissima (Strobl) Avon, Cambs., Somerset. iii-vi.

Genus Chonocephalus Wandolleck (Figs. 25, 28-30)

This genus includes about 40 species, mainly found in the tropics and sub-tropics. 3 species are recorded from the Palaearctic Region. The species reported from England has been transported around the world by man.

The larvae of a few species have been reported from rotting fruits, rotting palm spadices and similar situations.

20

9, With hair palisades on ventral edges of segments 1-3 of bind tarsi (fig. 30); tergite 3 of abdomen with 20-30 hilirs arising from pale spots embraced by a transverse dark band (fig. 29). CI'. Hypopygium as in fig. 28: wing as in fig. 25 .............. heymonsi Stobbe London (in hothouse). vi.

Genus Conicera Meigen (Figs. 10, 31-43)

This genus includes more than 20 species, of which 9 are known from Europe. The rest are from the Nearctic, Neotropical and Oriental Regions. Undescribed species are known from the Afrotropical Region.

C. tibialis is the infamous "Coffin Fly" that breeds in coffmed bodies that have been interred a year or more (Colyer. 1954a-c; Disney & Smith, 1978). It has also been reared from dead birds and other carrion. Other species are also associated with carrion. C. similis has also been reared from fungi. C. dauci larvae have been recorded in rotting plants. The adults visit flowers, particularly Umbelliferae.

Keys to Species Males

Claspers of hypopygium tapered to narrow points (figs. 31-32) ......... daucl (Meigen) Widespread in British Isles. iii-xi.

Claspers of hypopygium developed as irregular lobes which are not tapered and which tend to be rounded, even expanded, distally (figs. 33-37) ............................. 2

2 Posterior face of mid-femur with a sense organ consisting of a pit from which a furrow leads to an apical, hollow process (figs. 38-39) ...................................... 3

Posterior face of mid-femur without such a sense-organ. (To proceed the hypopygium must be mounted beneath a cover-slip in such a manner that when the latter is gently pressed down the right clasper is clearly displayed with the outer face uppermost - as figs 33-37) . ........................................................................ . 4

3 Pit of sense-organ on mid-femur larger and apical process longer (fig. 38). (Right clasper as fig. 33) .................................................... tibialis Schmitz Widespread in British Isles. iv-xi.

Pit of sense-organ on mid-femur smaller and apical process shorter (fig. 39). (Right clasper as fig. 34) ................................................... simHis (Haliday) Widespread in British Isles. iii-x.

4 The upper, inwardly-directed, lobe of right clasper of hypopygium rounded andonJy a little narrower than ventral lobe (fig. 35) .............................. tarsalis Schmitz Widespread in British Isles. iii-vii.

The upper lobe of right clasper of hypopygium a narrow process (figs 36-37) ......... 5 5 The ventral lobe of right clasper of hypopygium with 4-7 blunt, rounded, teeth in a single

row on the more-or-Iess straight inwardly-directed margin, and typically with a single pointed spine on lower edge near the lowest tooth (fig. 36) ..... scbnittmanni Schmitz Essex, Suffolk, Surrey, North Yorks. Ireland: Offaly. vi-ix.

The ventral lobe of right clasper with a dozen or more teeth in two irregular rows along the inside face of the inwardly-directed margin and extending around the lower edge (fig. 37). These teeth are somewhat tapered and are increasingly spine-like ventrally.floricola Schmitz Widespread in British Isles. iv-ix.

Females (For distribution and phenology see males)

Vein 3 without a small bristle at base, on upper side .............................. 2 Vein 3 with a small bristle at base on upper side ............................ , ..... 4

2 Hind tibia with 5-6 dorsal or antero-dorsal bristles, there being 2-3 bristles in the third quarter (fig. 40) .............................................. atypical simHis (Haliday)

Hind tibia with only 4 dorsal or antero-dorsal bristles, there being only 1 bristle in the third quarter (figs 41-42) .........••...............................•............. 3

3 Upper dorsal bristle of hind tibia always distinci:Iy above the upper antero-dorsal bristle and usually clearly shorter than bristle just below middle (fig. 41), ...•... daacl (Meigen)

Upper dorsal bristle of hind tibia level with or, more usually, below the upper antero-dorsal bristle and usually longer than bristle just below middle (fig. 42) ..........•........

21

..................................... typical simills (Haliday) or tibialis Scbmitz (l have, as yet, found no consistent difference between the females of these two species)

4 The hind tibia bristles of C. f1oricola (fig. 43) are very similar to those of typical C. similis and C. tibialis (fig. 42). Until females of C. tarsalis and C. schnittmanni caught in copula • or in reared series, are avallable it is not possible to separate the females of these two species from C. floricola.

Genus Diplonevra Lioy (Figs. 8, 46-60, Plates A and B)

This genus includes nearly 70 species and is known from all regions of the world. 24 species are known from the Palaearctic Region.

Some species breed in carrion. D./unebris has been reared from wasp (Vespula) nests. D. pilosella has been reared from a wounded earthworm. Adults of some species visit flowers, particularly Umbelliferae.

Keys to species Males

In most cases it is essential to remove a hind leg, with the trochanter intact, and to view it from the posterior face. 1 Hind tibia without antero-ventral bristles but with 2 or more antero-dorsais which are at least

as strong as the dorsal apical bristle .......................................... 2 Hind tibia with at least 1 (usually several) short antero-ventral bristles. Antero-dorsals, when

present, always weaker than dorsal apical bristle ............................... 3 2 Vein 4 straight (fig. 56). Basal half, or more, of hind femur yellowish. Posterior face of hind

femur at base and hind trochanter without blunt spines or stout bristles (fig. 46). Hind tibia with 3 hair palisades. Haltere pale yellow .............. abbreviata (von Roser) Widespread in S. of England. Cambs., Hereford. vi-x.

Vein 4 distinctly curved (figs. 8 and 57). Hind femur either uniformly dark brown or with basal half, or more, paler brown. Posterior face of base of hind femur with a variable, but highly characteristic row of blunt, tooth-like, spines, and hind trochanter with bristles and short, stout, spines (figs. 47 and 48). Hind tibia with 2 hair palisades. Haltere brown to dark brown ........ '" .................................... florea (Fabricius) Widespread in British Isles. iv-viii .

.3 Halteres yellow .............................................................. 4 Halteres black ............................................................... 5

4 Posterior face of base of hind femur with an ascending process bearing a single row of bristles along its entire length {fig. 49) ................................. uitidula (Meigen) Widespread in British Isles. iv-xi.

Base of hind femur with an ascending process bearing bristles at the tip only (fig. 52) .. 5 5 Proboscis elongated, narrow, and elbowed. Hind trochanter with cluster of short black 'studs'

in place of bristles (fig. 50) ................................... funebris (Meigen) Widespread in British Isles. vi-x.

proboscis short and frequently bulbous basally. Hind trochanter with normally tapered bristles ......................................................................... 6

6 Hind tibia with 1 or more short antero-dorsal bristles (when only 1 is present it is in the basal quarter). Base of posterior face of hind femur with characteristic cluster of bristles without an ascending process (fig. 51) ................................. concinna (Meigen) Widespread in British Isles. v-x.

Hind tibia usually devoid of antero-dorsal bristles, but a single small one is occasionally present. Base of hind femur with an ascending process, bearing 2-3 bristles (variable in both species). arising from rest of bristle cluster on posterior f~ (figs. 52 and 53) ......................................................................... 7

7 Right side of epandrium with only one bristle-like hair, the other hairs shorter than those on the ventral edge of anal tube and relatively sparse (jIg. 54). Anal tube typically projecting posteriorly in a more-or-Iess straight line that continue~ the line of the dorsal edge of

epandrium (fig. 54) ............................................ glabra Schmitz Widespread in England. Scotland: Nairn. iv-viii.

Right side of epandrium with several bristle-like hairs which are longer and more robust than those on the ventral edge of anal tube; these along with the shorter hairs are relatively dense at posterior margin (fig. 55). The anal tube typically tends to be somewhat pendulous, so it is clearly divergent from the line of the dorsal edge of epandrium (fig. 55) ........ . · ........................................................... pilosellaSchmitz Widespread in England and Scotland. iv-xi.

Females (For distribution and phenology see males)

Hind tibia without antero-ventral bristles but with 2 or more antero-dorsals which are at least as strong as the dorsal apical bristle .......................................... 2

Hind tibia with at least 1 (usually several) short antero-ventral bristles. Antero-dorsals, when present, always weaker than dorsal apical bristle ............................... 3

2 Vein 4 straight (fig. 56). Basal half, or more, of hind femur yellowish. Hind tibia with 3 hair palisades. Haltere pale yellow ...... , ., .................... abbreviata (von Roser)

Vein 4 distinctly curved (figs. 8 and 57). Hind femur either uniformly dark brown or with basal half, or more, paler brown. Hind tibia with 2 hair palisades. The abdomen may be extensively reddish. Haltere brown to dark brown ................ f10rea (Fabricius)

3 Pleural region of abdominal segment 6 with hairs around spiracle, these hairs being more than 3 x longer than the diameter of spiracle (fig. 58); a smaller patch of similar hairs on segment 5 behind the spiracle ................................. funebris (Meigen)

Apart from hairs at posterior border of segment 6 any hairs present on the pleural regions of abdominal segments 5 and 6 are less than 3 x length of diameter of spiracle (as on segment 4 of funebris, fig. 58) ...............................................•...... 4

4 Ventral face of each labellar lobe with less than 30 short, colourless, isolated spines in addition to longer hairs. Halteres usually yellowish ..................................... 5

Ventral face of each labellar lobe with more than 50 short, colourless, crowded spines in addition to longer hairs. The edge of this field of spines is clearly visible in prome when the lobe is viewed from the dorsal face (fig. 59). Halteres dark grey to black ....... 6

5 Microsculpture of posterior face of middle of hind femur is a mosaic.of simple polygons (fig. 60 and Plate A) .............................................. nitidula (Meigen)

Microsculpture of posterior face of middle of hind femur is a mosaic of polygons each of which tends to be made up of pigmented strips that are variously fused and frequently confluent with strips of adjacent polygons. The resulting polygons appear less sharply demarcated and have the appearance of being diagonally hatched. (Plate B) ........ . · ............................................................ glabra Schmitz

6 Hind tibia with 1 or more (usually 2 or 3) antero-dorsal bristles. Middle and hind femora more-or-Iess black with yellowish tips. Front coxae and femora dark brown. Wing brownish grey and somewhat darker beyond end of costa and narrowly along line of each thin vein · ......................................................... concinna (Meigen)

Hind tibia with 0-1 (usually 0) antero-dorsal bristles. Middle and hind femora brown with yellowish tips. Front coxae and femora brown, the latter with more than tips yellowish. Wings greyish to yellowish grey, not obviously darker beyond costa or along line of each thin

vein .......................................................... pllosella Schmitz

Genus Dohmiphora Dahl (Figs. 3 and 101)

This genus includes more than 100 species from all parts of the tropics and sub-tropics. The species now'well established in England has been transported around the world by man.

D. cornuta is a polyphagous saprophage in the larval stage, breeding in sewage, compost, dead insects, dead snails, rotting cargoes in ships (rice bran, cow peas, etc), vertebrate carrion and similar situations. It will also attack other iilsect eggs, larvae

23

and pupae (e.g. those of Psychoda altemata in trickling filter sewage beds) that it encounters competing for its food.

Hind tibia without bristles, apart from apical spurs. Legs yellow apart from some brown pigment on mid-coxae. 0", sensory complex on posterior face of base of hind femur as in fig. 101 (note also the few scattered hairs, near the dorsal edge - these hairs form a dense cluster in the closely related afrotropical D. fraudans, and are missing altogether in other species with a similar sensory complex). Halteres yellow. Q, without tergites on abdominal segments 5 and 6 ................................... cornuta (Bigot) Sporadically occuring in Berks, E. Anglia, Midlands, N. Yorks, N. Wales. iii-x.

Genus Gymnophora Macquart (Figs. 61-74)

This genus includes about two dozen species in the Palaearctic, Oriental, Nearctic and Neotropical Regions. 6 species are known from Europe.

The larval natural history is unknown.

Keys to Species Males

The hypopygium is often withdrawn. It can be expelled by judicious pressure on the coverslip during mounting.

Oblique ridge of notopleuron (fig. 65) with posterior border black, in contrast to its fringe of pale hair· .............................................................. 2

Oblique ridge of notopleuron largely pale. Hypopygium as fig. 61. Small swelling of costa before tip of vein 1 embracing a pale oval spot . . . . . . . . . . . . . . . . .. arcuata (Meigen) Widespread in British Isles. vi-ix.

2 Venter of abdominal segment 6 usually with relatively long hairs (at least in the posterior row) and these hairs clearly in more than one row. Hypopygium with relatively short cerci and as fig. 62 ........•.................................. quartomollis Schmitz Widespread in Great Britain. vi-viii.

Venter of abdominal segment 6 either with a single row of hairs or the hairs are clearly shorter (i.e. clearly shorter than those on cerci). Cerci relatively long (figs. 63-64) ......... 3

3 Swelling of costa before tip of vein 1 without a distinct pale spot (fig. 69). Hypopygium as fig. 63 ...................................................... healeyae Disney Cumbria, Durham, N. Yorks., Suffolk. vi-ix.

Swelling of costa before tip of vein I embracing a distinct, pale, oval spot. Hypopygium as fig. 64 . . . . ........... _ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. integralis Schmitz Gloues. x.

Females (For distribution and phenology see males)

Oblique ridge of notopleuron (fig. 65) with posterior border black, in contrast to its fringe of pale hair ............................................................... 2

Oblique ridge of notopleuron largely pale. Swelling of costa before tip of vein 1 embracing a pale oval spot. Abdomen with reduced tergite on segment 3 and with vestigial tergites only on segments 4 and 5 (unless critically examined they appear to be absent altogether) ........................................................... arcuata (Meigen)

2 Abdominal tergite 3 reduced, tergite 4 missing, tergite 5 present and well developed, tergite

24

6 narrows posteriorly (fig. 70). Internal paired glands opening between tergites 5 and 6 obscure (critical illumination is required in order to reveal them at all) (fig. 70). Tergite 8 (when exposed to view) as in fig. 66 or a little narrower in some specimens. Swelling of costa before tip of vein 1 slight and without a pale oval spot .... quartomollis Schrnitz

Internal paired glands opening just before anterior margin of tergite 6 distinct (figs. 71 and

73). Tergite 5 missing (fig. 71) or if present the common opening to the paired glands is very distinctive and conspicuous (fig. 73) ...................................... 3

3 Internal paired glands as fig. 71. Abdominal tergite 7 sparsely haired (fig. 72). Tergite 8 (when exposed to view) as in fig. 67. Swelling of costa before tip of vein I with no pale spot, or if present, it is obscure (fig. 69) ................................. bealeyae Disney

Internal glands as in fig. 73, discharging into a common chamber whose opening to the exterior is transversely elongated and whose margin is conspicuous (fig. 73). Tergite 6 with anterior part almost separated from larger posterior part by a 'waist' (fig. 73). Tergite 7 with long conspicuous hairs (fig. 74). Tergite 8 (when exposed to view) hairless and as fig. 68. Swelling of costa before tip of vein 1 embracing a distinct, pale, oval spot. .. integralis Schmitz

Genus Gymnoptera Lioy Figs. 44-45)

There are 6 species known in this genus, from the Oriental, Neotropical and Palaearctic Regions. The 2 Palaearctic species both occur in Britain.

G. /ongicostaiis was only distinguished from G. vitripennis in 1933. Failure to accommodate this fact has led to much uncritical citation of observations published prior to 1933. G. longicostalis normally breeds in bumble-bee nests (Bombus terrestris and B. lucorum being recorded). G. vitripennis normally breeds in wasp nests (Vespula germanica and V. vulgaris being recorded). Both species have also been reared from the same caterpillar of the Carpenter Moth (Cossus cossus).

Key to Species Males and females

CJ' , posterior face of base of hind femur with several furrows, which are more divergent from ventral edge of femur (fig. 44). 9, a short, but distinctly differentiated, spur situated dorsally or antero-dorsally on front metatarsus, on the second tarsal segment. and usually on the third segment as well. 2, or more, costal cilia of dorsal row situated beyond junction of vein I ............................................... .IongicostaJis Schmitz Widespread in lowlands of British Isles. vi-vii, x.

CJ', posterior face of base of hind femur with fewer furrows, which are less divergent from ventral edge of femur (fig. 45). 9, a short dorsal or antero-dorsal spur present only on metatarsus of front leg. At most a single costal cilium of dorsal row situated beyond junction of vein 1. ................................................ vitripennis (Meigen) Widespread in lowlands of England. viii-xi.

Genus Hypocera Lioy (Fig. 75)

There are 6 species known in this genus, from the Oriental, Nearctic and Palaearctic Regions. The single Palaearctic species occurs in Britain.

The larval natural history is unkown. Dark brown to black species with robust bristles on frons. 2 anterior bristles on hind tibia

(fig. 75). Wing membrane dusky grey, but yellowish at base. CJ'. hypopygium with numerous long hairs on epandrium and anal tube. 9, with proboscis somewhat elongate and protruding from beneath the projecting fronto-c1ypeus .......... mordellaria (Fallen)

E. Anglia, Hants., Hereford., O;iford., N. Wales. vi"vii.

Genus Metopina Macquart (Figs. 76-96)

This genus includes more than 30 species from all regions of the world. 10 species

25

are known from Europe. The larval natural history is unknown. However there is a specimen of M. oligoneura

in the Hope Entomological Collections (University Museum, Oxford) with the label "fr. Phorbia debris" .

The adults visit flowers. A continental species has been observed persuading ants to regurgitate food on which it then fed.

Keys to Species Males

It is necessary, in most cases, to mount a hind leg on a slide with the posterior face uppermost and the trochanter intact. Likewise the abdomen should be detached as close to the thorax as possible, mounted ventral side up and gentle pressure applied to the coverslip in order to display the mid-venter clearly. 1 Costa unusually thickened, with maximum width clearly wider than subcostal cell and wing

generally larger and broader basally than usual (fig. 77) ......... crassinervis Schmitz Berkshire vii.

Costa normal, wing of normal size and narrower basally (e.g. fig. 78) ............... 2 2 Ventral face of abdominal segments 3-6 with (). 7 hairs. When present these hairs are as large

as, or larger than, those on tergites and are either restricted to a transverse row on segment 6 or they form an irregular median row on segments 3-6 ......................... 3

More than 7 hairs are present in a distinct patch on the ventral face of segments 3-4. In some cases these hairs are situated on a distinctly chitinised platelet (figs. 79-80). An irregular transverse row of hairs on segment 5 and usually a few hairs on segment 6 ......... 4

3 Posterior face of hind femur with 8-14 furrows running in an inclined transverse band just before the half-way point (fig. 83) ................................. ulricbi Disney Avon, Bucks., Essex, Salop., Surrey, Sussex, Wilts., vi-viii.

Hind femur without this patch of furrows ......................... galeata (HaJiday) Widespread in England, S. Wales, Ireland: Galway, Offaly. vi-x.

4 Posterior face of base of hind femur with conspicuous "sensory patch" that includes a distinct circular pit (fig. 84). Hairs of hind trochanter reduced in number and with only a single, basal one evident on ventral face, which is itself fringed with dense microscopic hairs. In addition the terminal spine is dark, somewhat reduced and strongly tapered (fig. 84). (Abdominal venter with a distinct, but pale, ventral plate on segment 4. This plate bears 15-25 hairs, some being on the disc) ........................... heselhausi Schmitz Hants., Surrey, Wilts. iv, viii.

Sensory patch of hind femur less conspicious and without a pit (figs. 85-88). At least 3 hairs along ventral face of hind trochanter, which is itself fringed with sparse, inconspicuous microscopic hairs. The terminal spine is more robust, pale, and more gradually tapered (figs. 85-88) ................................. , ............................. 5

5 Venter with a distinct ventral plate on segment 4, bearing irregular rows of hairs along the lateral margins but none along the median third (fig. 80). The irregular extension of this plate anteriorly bears some hairs in a more median position. The main plate usually has an obviously concave posterior margin (fig. 80). Sensory patch on posterior face of base of hind femur usually visible as a darker smudge at relatively low magnification (fig. 85) .......................................................... . oHgoneura (Mik) Widespread in British Isles. v-xi.

Venter without a clearly defined plate on segment 4, although there may be an ill-defined, incipient plate developed from areas of pigmentation extending from the base of each hair (fig. 81). Sensory patch at base of hind femur obscure (figs. 86-88) ............... 6

6 Venter with hair patch on segment 4 with less than 16 hairs arranged in two irregular series

26

more-or-Iess separated by a bare median band (fig. 81). The areas of pigmentation spreading from the base of each hair usually show some coalescence, thus forming an incipient plate (fig. 81). Micro-sculpture at base of posterior face of hind femur with long axes of polygons subparallel to ventral edge of femur (fig. 86) ...................... pileata Schmitz Widespread in S. of England. E. Anglia. S. Midlands, Powys. vi-bc.

Venter with 12-30 hairs not separated into two irregular series and areas of pigment at base

of hairs obscure and showing no indication of coalescence (fig. 82). Microsculpture at base of posterior face of hind femur with long axes of polygons mainly divergent from ventral edge of femur (figs. 87-88) .................................................. 7

7 Microsculpture of posterior face of hind femur with more-or-Iess straight rows of polygons (fig. 87). Venter with hair patch on segment 4 usually with less than 20 hairs ....... . · ............................................................ braneri (Strobl) Cambs., Norfolk, Oxford, Surrey, Scotland: Argyll, Ross. v-vii.

Microsculpture of posterior face of hind femur with rows of polygons curving dorsally (fig. 88). Venter with hair patch on segment 4 usually with more than 20 hairs ....... , .. . · ............................................................ perpnsilla (Six) Widespread in S. England and E. Anglia, Cheshire. v-x.

Females (For distribution and phenology see males)

When slide-mounting the abdomen should be detached as close to the thorax as possible. It should be mounted dorsal side up in such a manner that tergites 4 and 5 are displayed clearly. 1 Fourth abdominal tergite at least twice width of fifth tergite (fig. 89). Typically there is a

dimple-like hollow, demarcated by microscopic denticles, either side of the fourth tergite. The pleural region of segment 5 may be inflated to produce a hemispherical protuberance each side ................................................... galeata (Haliday)

Fourth tergite approximately same width as fifth tergite or only a little wider (figs. 90-96) .................................................. , ...................... 2

2 Anterior flap of fifth tergite longer than broad, and tapered posteriorly (fig. 90) ...... . · ............................................................ braneri (Strobl)

Anterior flap of fifth tergite not thus (figs. 91-96) ................................ 3 3 Anterior flap of fifth tergite clearly more than a semi-circle, with the widest point obviously

posterior to the anterior margin (figs. 91-92) ....... , .............. , .. , ......... 4 Anterior flap of fifth tergite more-or-Iess semi-circular and widest point at or near anterior

margin (figs. 93-96) ........................................................ 5 4 Fourth tergite narrower and with hairs of posterior margin markedly longer than those on

disc (fig. 91). Costa not conspicuously thickened, but with maximum width clearly less than maximum width of subcostal cell. Wing of normal size and shape (cf. fig. 78) ... · . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ulrichi Disney

Fourth tergite very much broader than long and hairs on posterior margin and disc of appproximately equal size (fig. 92). Costa conspicuously thickened, so that the maximum width is clearly greater than maximum width of subcostal cell. Wing larger and broader basally than usual (fig. 76) .................................. crassiDervis Schmitz

5 Hairs of abdominal venter in narrow band, there being less than 12 in the posterior transverse row on segment 4. Anterior flap of tergite 5 without a regular semi-circular ouiline, but with distinct sides (which may be more-or-less subparallel) and posterior border (fig. 93) · ............................................................ perpnsilla (Six)

Hairs of venter in broader band on segment 4 and 5, there being more than 15 in the posterior transverse row on segment 4. Anterior flap of fifth tergite is a more-or-less regular semi-circle in outline (figs. 94-96) ................................................. 6

6 Swarms of hairs of dorsal and ventral sides of abdominal segments 4 and 5 linked only by a single row over the pleural region of segment 5 and even this row of hairs may be interrupted. Tergite 4 not wider than fifth and usually with an unpigmented median band extending approximately half way from anterior margin (fig. 94) ..... pUeata Schmitz

Swarms of hairs of dorsal and ventral sides of abdominal segments 4 and 5 more-or-Iess linked by at least 2 rows extending over the pleural region on segment 5 and 1 row on segment 4. Tergite 4 slightly wider than 5 (fIgs. 95-96) and never with more than a slight emargination of the median anterior border.- .....................................•........ 7

7 Abdominal tergites 4 and 5 relatively small (rig. 96) (4 is less than 0.2 mm wide). The posterior part of 5, while it is a little variable (being sometimes more, sometimes less, tapered than as illustrated - fig. 96), has narrower anterior arms and its total area relative to that of the anterior, semi-circular, flap is clearly less ..................... oUgonenra (Mik)

27

Abdominal tergites 4 and 5 relatively large (fig 95) (4 is nearly 0.3 mm wide). The posterior part of 5 has broader anterior arms and its total area relative to that of the anterior flap is clearly more ............................................. heselhausi Schmitz

Genus Phalacrotophora Enderlein (Figs 97-100)

This genus includes .about 50 species from all regions of the world. 6 species are known from Europe.

The larvae of the British species parasitize the pupae ofladybird beetles (Coccinellidae)

Key to Species Males and females

Hind metatarsus usually somewhat swollen and usually dark brown to black, C1', witb rigbt lobe ofbypandrium elongated so tbat is projects to tip, or beyond end, of epandrium (fig. 98). 9, witb end of ovipositor simple and sbort-haired (fig. 100) .... fasciata (Fallen) Widespread in S. England and E. Anglia, Salop. iii-viii.

Hind metatarsus at most slightly swollen and yellow to yellow-brown in colour. C1', without elongated lobe on right side of hypandrium (fig. 97). 9, with end of ovipositor bearing a pair of sub-terminal, dorsally projecting, tborn-like processes, and ventrally bearing long hairs (fig. 99) ............................................. berolinensis Schmitz Widespread in lowlands of England and Wales. Scotland: Perths., Ireland: Offaly. v-vii.

Genus Phora Latreille (Figs. 102-126, 181-182)

This genus includes at least three dozen species in the Afrotropical, Nearctic, Neotropical, Oriental and Palaearctic Regions. At least two dozen occur in Europe.

The larvae of P. holosericea live 20-40 cm below the soil surface preying upon root ap!}ids.

Recognition of species in this genus has been subject to a high frequency of error due to over-reliance on bristle and wing characters which have been found to be much more variable than supposed by previous workers. For this reason species recognition is now firmly based on details of the male hypopygium. In view of the frequency of misidentifications encountered when examining museum collections it will not be possible to provide a key to females until specimens caught in copula with certainly identified males are available for all species (or else reared series of males and females). The author still lacks such females for P. artifrons, P. bullata, P. edentata, P. hamata, P. obscura, P. praepandens and P. speighti. P. bullata was only added after the typescript had been prepared, hence the figures are out of place at the end!

Key to Species Males

It is essential to slide-mount the hypopygium. The entire hypopygial capsule is readily teased out from the rest of the abdomen. It should be mounted on its side with the left side facing up. The right side can be examined by turning the slide over, supporting it at either end to prevent fouling of the mount (e.g. by a slide placed either side of the microscope stage). It should be noted that the posterior process of the left side of hypandrium is easily rotated and so may be directed inwards towards the mid-line or downwards - it may, therefore, appear somewhat other than as drawn in the figures.

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Apart from the row of conspicuous dorsal bristles on the mid-tibia there are 2 anterior bristles in the upper half, although one may be only half the size of the other. The hind tibia often with 2 anterior bristles in upper half also ...................................... 2

Mid-tibia with only 1 anterior bristle in upper half. Hind tibia with only 1 bristle in upper half .......................................................................... 6

2 Left side of epandrium deeply cleft to give a shorter upper lobe and a longer lower lobe (figs 102 and 104) .............................................................. 3

Left side of epandrium not deeply cleft to give 2 distinct lobes (e.g.fig. 116) .......... 5 3 Frons conspicuously narrowed in dorsal half (fig. 124). Hind tibia with only 1 anterior bristle

in upper half. Hypopygium as figs. 104 and 105 .................. artifrons Schmitz Oxford. Scotland: Inverness. Sutherland. Wigtown. iv-vii.

Frons at most slightly narrowed in dorsal half (similar to fig. 126) ................. .4 4 Hind tibia with 1 or 2 (usually 2) bristles in upper half. Hypopygium as figs. 102 and 103

.......................................................... dubia (Zetterstedt) Widespread in North of England and Scotland. Ireland: Kerry, Offaly. iv-vi.

Hind tibia with only 1 bristle in upper half. Hypopygium otherwise ................. 6 5 Appendage of right side of epandrium expanding posteriorly to a strongly convex hind margin

(fig. 117). Hypandrium as figs. 116-117 ................. , ......... stictica Meigen Widespread in N. England, N. Wales and Scotland. vii-x.

Appendage of right side of epandrium bilobed so that it has a concave hind margin (fig. 123). Hypandrium as figs. 122-126 ................................................ 6

6 Left side of epandrium deeply cleft to give a shorter upper lobe and a longer lower lobe (figs 108, 1l0, 112 and 114). The upper lobe is sometimes abbreviated (fig. 181) ......... 7

Left side of epandrium not cleft in this manner, although the distal portion may be somewhat excavated or (in P. speighll) a lower lobe may be present but be concealed by the expanded upper lobe (figs 106, 116, 118, 120 and 122) .................................. 12

7 Right side of hypandrium with a black, relatively narrow, unturned process which is strongly tapered and almost thorn-like (fig. 109). Upper lobe of left side of epandrium with a somewhat concave distal margin (may not be obvious if not viewed from correct angle) and lower lobe of characteristic shape (fig. 108) ..................... am (Meigen) Widespread in British Isles. iv-xi.

Upturned process of right side of hypandrium brown to dark brown, broader, and less strongly tapered or not tapered at all (figs. 105, lII, 113, 115 and 182) .................... 8

8 Frons conspicuously narrowed in dorsal half (fig. 124) (aberrant specimens with only I bristle on mid-tibia) ...............................................•..... return to 3

Frons at most only slightly narrowed in dorsal half (similar to fig. 126) .............. 9 9 Upper lobe of left side of epandrium with a nearly vertical posterior margin which is irregularly

straight-edged to somewhat concave in middle, and is distinctly crenellate (fig. 110). Hypandrium as figs. 110-111. .............................. bolosericera Schmitz Widespread in British Isles. v-viii.

Upper lobe of left side of epandrium with curved or inclined posterior margin which is strongly convex posteriorly or postero-ventrally and is not crenellate (figs 112 and 114) ..... 10

10 Upper left lobe of epandrium relatively short, extending at most to level of tip of anal tube (fig. 181). Lower lobe with postero-ventral margin curving inwards (and folded forwards) beyond heavily sclerotised region (fig. 182); and postero-dorsally weakly sclerotised so that it includes translucent 'windows' (fig. 181). Appendage of right side of epandrium more dorsal in position and with expanded, somewhat 'warped' distal extremity (fig. 182). Upturned process of right side of hypandrium relatively large and broad, with an irregular (almost crenellate) anterior (upper) margin (fig. 182) ............... bullata Schmitz Suffolk viii.

Upper left lobe of epandrium longer, reaching well beyond tip of anal tube (figs. 112 and 114). Appendage of right side of epandrium more lateral in position (figs 113 and 115). Upturned process of right side ofhypandrium smaller and less robust (figs. 113 and 115) ........................................................................ 11

11 Upper left lobe of epandrium of smooth appearance with few, if any, wrinkles. Posterior margin of lower lobe without fringe of minute hairs (not to be confused with short hairs in upper halt) - fig. 112. Hypandrium as figs 112-113. Wings with membrane more-or-less clear .................................................... edentata Schmitz Widespread in lowlands of England, Wales and Ireland, v-ix.

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Upper left lobe of epandrium with rough appearance due to profusion of fine wrinkles. Posterior margin of lower lobe with fringe of minute hairs (in addition to short hairs in upper half) - fig. 114. Hypandrium as figs 114-115. Wings with membrane tinged grey to yellowish-brown ............................................. Uuctll Schmitz Widespread in British Isles. ii, v-lx.

12 Frons somewhat narrowed in dorsal half (fig. 125). Appendage of right side of epandrium relatively small (fig. 119). Hypandrium as figs 118-119 ......... praepandells Schmitz Scotland: Perths. vi.

Frons more or less parallel-sided (e.g. fig. 126). Appendage of right side of epandrium relatively large (figs 107, 121 and 123). Hypandrium otherwise ........................... 13

13 Appendage of right side of epandrium with strongly convex posterior (distal) margin (fig. 107). Left side of epandrium relatively simple (fig. 106). Upturned process of right side of hypandrium more sharply angled at point of upturn of ventral margin ~fig .. 10~) ... · ............................................................ spelgbti Disney Oxford. Ireland: Dublin. vi-vii.

Appendage of right side of epandrium with indented or strongly concave posterior (distal) margin (figs. 121 and 123). Left side of epandrium more complex (figs. 120 and 122). Upturned process of right side of hypandrium less sharply angled at point of upturn of ventral margin (figs. 121 and 123) ........................................... 14

14 Appendage of right side of epandrium with more deeply notched posterior margin (fig. 121). Process ofleft side of hypandrium with a relatively long basal process (as well as a shorter one) - figs 120 and 121. ...................................... hamata Schmitz Cambs. v.

Appendage of right side of epandrium with less deeply notched posterior margin (fig. 123). Process of left side of hypandrium with a shorter basal process (fig. 122) .......... . · ....................................................... obscura (Zetterstedt) Scotland: Inverness. v.

Genus Plectanocnema Schmitz (Figs. 26-27)

Only 1 species is known in this genus. Its natural history is unknown. Dark brown to blackish species with dark brown legs and halteres. Palps brown with numerous

short bristles along ventral face. Mesopleuron bare. Scutellum with 4 bristles. Wings brownish grey. (1, with posterior ventral corners of epandrium developed as rounded lobes extending as far as, or beyond, posterior extremity of dark-brown anal tube (fig. 27). · ........................................•.................. nudipes (Becker) Avon, Hereford, Kent, Somerset, Suffolk. iv-vi.

Genus Pseudacteon Coquillet (Fig. 12)

This genus includes more than 40 species from all parts of the world except the Afrotropical Region, where it is represented by the closely related genus Microselia. 6 species are known from Europe.

The larvae are parasitoids of adult worker ants. P. brevicauda parasitises Myrmica (M. ruginodis andM. scabrinodis). P.jormicarum attacks Lasius (L. flava and L. niger and Formica sanguinea, and possibly other genera (the possibility of other species of Pseudacteon being confused with P. jormicarum renders early records of other ant hosts a little suspect.) Specimens caught attacking ants should be preserved with some of the ants and the latter identified as well as the flies.

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Key to Species Males and females

Halteres with at least the tip of knob yellowish ................................... 2 Halteres completely dark ................ species not previously recorded from Britain

(see Schmitz, 1938a). 2 Mid-tibia without a dorsal hair palisade. 9 , stemite on segment 6 (adjacent to base of down-

curved ovipositor) with hairs only the longest being less robust than the longest hairs (as opposed to bristles) on thoracic scutum. (J', with palps and halteres largely yellowish ............................................ : ........... formicamm (Verrall) Widespread in Southern half of England. ii, vi-viii.

Mid-tibia with a dorsal hair palisade in basal half. 9, sternite on segment 6 with a row of bristles, which are clearly more robust than longest hairs on thoracic scutum. (J', palps brownish. Halteres with only distal half of knob pale ........... brevlcauda Schmitz Berks., E.Ang/ia, Kent, Hants., Surrey. vi-viii.

Genus Puliciphora Dahl (Figs. 127-129)

This genus includes more than 50 species from all regions of the world. Two species are known from Europe, but P. borinquenensis is really a tropical species that has been transported around the world by man.

P. borinquenensis breeds in organic refuse. The wingless female is transported by the male during mating.

9 , with anterior, semi-circular, flap of abdominal tergite 5 more than half width of latter (fig. 128). Frons with 4 supra-antenna! bristles, a pair of antero-laterals, a pair of pre­ocellars, a pair of ocellar bristles, and 4 bristles on vertex. Hairs of frons all simple (unlike the closely related P. rujipes in which the hairs either side of the ocellar triangle end in comb-like tridents). (J' , with hypopygium as fig. 129, and wing as fig. 127. Scutellum with 4 bristles .............................................. borinquenensis Wheeler Oxford. (As pest in laboratory insect cultures).

Genus Spiniphora Malloch (Figs. 11, 130-133)

This genus includes about 20 species from the Nearctic, Oriental and Palaearctic Regions. 9 species are known from Europe.

The larvae live in dead snails, but have also been recorded from other situations (e.g. S. bergenstammi has been reared from an old nest of a blackbird; and is frequently reported by public health authorities from improperly washed milk bottles).

Keys to Species Males

Front and middle legs dark brown. A dark smudge usually evident around origin of vein 4 .......................................................... macniata (Meigen) Widespread in England. Anglesey. Ireland. iii-v, viii.

Front and middle legs yellow to yellowish-brown. No dark smudge around origin of vein 4 ......................................................................... 2

2 Hind tibia with 2 bristles in upper two-thirds (an upper antero-dorsal and a lower antero-ventral) ..................................................... dorsaDs (Beeker) S. W.England. W.Midlands. S. Wales. S. W.Scotland. Ireland: Mayo v-vii,x.

Hind n"ia with 3 bristles in upper two-thirds (an upper antero-dorsal, a lower antero-ventral, and a true dorsal near or below the latter) ..................................... 3

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3 Palp large relative to segment 3 of antenna, with relatively short bristles, and with no trace of an irregular indentation (fig. 130) .............................. excisa (Becker) Ireland: Offaly. Wicklow. vii.

Palp relatively small, with longer bristles and with an irregular indentation on outer face (fig. 132) ..................................................... bergenstammi (Mik) Widespread in England. Scotland: Dunbarton. Ireland. v-ix.

Females (For distribution and phenology see males)

Front and middle legs dark brown. A dark smudge usually evident around origin of vein 4 · ......................................................... maculata (Meigen)

Front and middle legs yellow to yellowish-brown. No dark smudge around origin of vein 4 ..................................•.......••....•........................ 2

2 Hind tibia with 2 large bristles in upper two-thirds (an antero-dorsal above and an antero-ventral below) plus a dozen, or more, shorter dorsal bristles along length of tibia .... · ........................................................... dorsalis (Beeker)

Hind tibia with 3 large bristles in upper two-thirds (an upper antero-dorsal, a lower antero­ventral, and a true dorsal near or below the latter) but without a series of shorter dorsals ......................................................................... 3

3 Palp entire, with no trace of an irregular indentation (fig. 131) ......... excisa (Becker) Palp with an irregular indentation on outer face (fig. 133) ......... bergenstammi (Mik)

Genus Triphleba Rondani (Figs 6, 134-178, 183-186)

This genus includes about 90 species, from the Palaearctic and Nearctic Regions. Burma and New Zealand. About 55 species are known from Europe.

Several species are known to breed in vertebrate carrion. T. antricola is known to breed in bat dung in caves (but the species is not confined to caves). T. lugubris breeds in nests of wasps (Vespu[a spp.) T. minuta breeds in the fungus Gymnopilusjunonius ( = Pholiota spectabilis). Adults of some species visit flowers, particularly Umbelliferae.

T. subcompleta was only added after submission of the typescript (Disney, 1983). It has been necessary to insert the relevant figures out of sequence at the end.

Keys to Species Males

Vein 3 unforked or with fork incomplete (the base of vein 2 being absent, so that vein 2 is not attached to, and originating from, vein 3 - e.g. fig. 134) .......... ~ ......... 2

Vein 3 forked (i.e. vein 2 is not only present but it arises from vein 3) ............... 6 2 Segment 3 of antenna drawn out into a conspicuous point so that its maximum length is more

than 1.5 x maximum breadth (fig. 135). Process of left side of upper half of epandrium narrows distally and curves in towards mid-line (fig. 140). Vein 2 completely absent · ....................................................... citreiformis (Becker) Widespread in N.England and Scotland. Cambs., Hereford. Ireland: Galway. vii-x.

Segment 3 of antenna not drawn out in this manner and if the length is more than 1.5 x maximum breadth the tip is truncate (fig. 136) ................................. 3

3 Segment 3 of antenna long haired and drawn out to a truncated point (fig. 136). Legs largely yellowish. Halteres yellow. (processes of upper half of epandrium subequal in length. Distal half, or more of vein 2 present) ............................... mianta (Fabricius) Widespread in lowlands of England, Scotland: Dunbarton. ix-x.

Segment 30r antenna slightly pointed (fig. 137) or more-or-less rounded (fig. 138). At least middle and hind legs largely brown. Halteres with at least knob largely brown or blackish •.......••.•.•...•...••..........•........•...........•....•.•.•......... 4

4 Fore coxae yellowish, contrasting with brown femora. Wing membrane greyish with brownish (or dark greyish) tint in places, particularly beyond end of costa and bordering veins 4-6.

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The latter veins brown and readily discerned at low magnifications (c.g. x 10). (Proccsses of upper half of epandrium subequal in length and conspicuously haired, particularly the one on the right side. Vem 2 varies from being totally absent to being complete (but normally only the distal half is present).) ................................... gracllis (Wood) Essex. Hereford., Widespread in N.England. Scotland: Dunbarton. Ireland. viii-xi.

Fore coxae brown and not contrasting in colour with femora. Wing membrane greyish without brown (or darker greyish) tint. Veins 4-6 pale and not easily discerned at low magnifications ......................................................................... 5

5 Vem 2 completely absent. Process of upper half of left side of epandrium more-or-Iess parallel-sided until just before the broadly rounded tip, which is devoid of hairs (fig. 141.) Process of right side distinctly shorter. Segment 3 of antenna somewhat pointed (fig. 137) .... · ............................................................. smitbi Disney Hereford. viii.

At least tip of vein 2 usually discernible (fig. 134). Process of upper half of left side of epandrium tapers to a narrowly-rounded tip, which is over-reached by some of the hairs (fig. 142). Process of right side subequal in length to that ofleft side (figs. 143-144). Segment 3 of antenna more rounded (fig. 138) ......................... crassinervis (Strobl) Durham. Gwent. ix-x.

6 Hairs of arista (in distal half) shorter than maximum breadth of base of segment 3 of arista ...................•....................••............................... 7

Hairs of arista (in distal half) longer than maximum width of base of segment 3 of arista ......................................................................... 8

7 Processes of upper half of epandrium with that of left side a little longer, clearly broader, and somewhat square-ended compared with that of right side (figs 183-184). Hair at base of vein 3 approximately equal in length to costal cilia. Axillary ridge of wing with 1-3 (usually 2) bristles .................................................. Iugubris (Meigen) Widespread in British Isles. v-viii.

Processes of upper half of epandrium sub-equal in length, with that of left side broader basally than that of right but tapering as the latter (figs. 185-186). Hair at base of vein 3 clearly longer (at least 1.3x) than costal cilia. Only 1 bristle on axillary ridge of wing ....... . · ....................................................... subcompleta Schmitz Scotland: Renfrew. vii

8 Scutellum with a posterior pair of bristles and an anterior pair of short fine hairs (clearly shorter and finer than hairs of scutum). Halteres variable in colour ............... 9

Scutellum with 2 pairs of bristles, both of which are clearly more robust and longer than hairs of scutum. Halteres always dark brown to black .............................. 20

9 Halteres yellowish to pale yellow .............................................. 10 Halteres brownish to black .................................................... 15

10 Legs largely yellowish apart from tendency to somewhat brownish femora. (Hypopygium as fig. 150) .............................................. luteifemorata (Wood) Widespread in England. viii-xi.

Legs largely dark in colour ............... ; ................................... 11 11 Palps swollen, being at least as wide as antennal segment 3, and with only a few, short, bristles

apart from a single, longer terminal one (fig. 6). Fork of vein 3 relatively small and with vein 2 relatively long (fig. 145) ............................... nudipalpis (Becker) Widspread in British Isles. iii-xi.

Palps never as wide as antenna! segment 3 but if, never-the-less, a little swollen then the fork of vein 3 is relatively large and with vein 2 relatively short (fig. 146) .............. 12

12 Ventral edge of hind femur with dense patch of short hairs, most of which have their tips curled over (fig. 139) ........................................ excisa (Lundbeck) Scotland: Moray. xi

Ventral edge of base of hind femur with less crowded hairs, which are similar to those on the anterior face .......................................................... 13

13 Vein 3 with unusually large fork and with 1-5 halrs on dorsal face before fork, these hairs being shorter and fmer than bristle-like hair at base of vein (fig. 147) .............. . · ........................................................ autumnalis (Beeker) Cambs., Herts., Suffolk. i-ii,ix.

Vein 3 with smaller (fig. 149) or narrower (fig. 146) fork and without fine hairs on dorsal face (apart from the single, bristle-like hair at base) ............................ 14

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14 Upper process of right side of epandrium subequal in length to process of left side, but much narrower than latter (figs 151-152) ............ " .............. hyalinata (Meigen) Essex, Hereford., Herts., Oxford, Sussex. Scotland: Ross and Cromarty.. Ire/and. i-v.

Upper process of right side of epandrium very much shorter than process of left side (figs 153-154) ...................... : ......................... .intempesta (Schmitz) Cumbria, G/oucs., Hants., Surrey. Ireland: Offaly. ii-iv, xi-xii.

15 Wing membrane greyish with brownish (or darker greyish) tint in places, particularly beyond end of costa and bordering veins 4-6. Fore coxae yellowish, contrasting with brown femora. (see couplet 4) ................................................ gracilis (Wood)

Wing membrane paler grey or more-or-Iess clear. and without darker tint bordering veins 4-6. Fore coxae same colour as femora, although they may be a little paler distally ... .......................•......... ............... ...................•... . 16

16 Processes of epandrium subequal in length ..................................... 17 Process of right side of epandrium very much shorter than process of left side (figs 155 and

157) .................................................................... 18 17 All femora brownish or brownish grey. Palps dusky yellow to darker. Vein 3 either uniformly

brown or median third or less (running length of vein) is paler. Even this paler stripe is still darker than wing membrane. The longest hairs of cerci usually over-reach extremities of processes of epandrium ................................... antrlcola (Schmitz) Devon, Hereford., N. Yorks. Scot/and: Wigtow~. Ireland: Kerry, Wicklow. iii-x.

All femora in part at least dusky yellow, although the hind femora may be largely greyish. Palps clear yellow to slightly dusky. The median third, or more, of vein 3 is as pale as membrane of wing. The longest hairs of cerci usually end at or before level of extremities of processes of epandrium ......................•.......... distinguenda (Strobl) Widespread in British Isles. v-x.

18 Palps exceptionally large and with a few small hairs only (fig. 156). Hypopygium as figs 155 and 157, with. shovel-shaped process of left side of epandrium .... flexipalpis Schmitz Hereford. x.

Palps not obviously enlarged, and bearing distinct bristles ........................ 19 19 At least the front legs yellowish. Process of left side of epandrium strongly tapered (fig. 150)

{see couplet 10) .......................................... Iuteifemorata (Wood) Legs brown to dark grey. Process ofteft side of epandrium as fig. 158 (it may appear a little

tapered if viewed not directly from the side) ........................ vitrea (Wood) Cambs., Hereford., N. Yorks. ix-x.

20 Left side of epandrium with process divided into two arms (figs 159-160) ........... 21 Process of left side of epandrium not divided (figs 161, 164-165) ................... 22

21 Both arms of process ofteft side of epandrium narrow and as in fig. 159. There is also a median process extending dorsally and posteriorly between the normal lateral processes of epandrium (fig. 159). Hind tibia usually without a bristle in upper half ............ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . opaca (Meigen) Widespread in England and Scotland. ii-v.

Lower arm of process ofteft side of epandrium very much larger than upper arm (fig. 160). Hind tibia usually with a bristle in upper half on anterior face .. intermedia (Malloch) Widespread in England. Scotland: Dunbarton. i-vi.

22 Process of left side of epandrium short but with a cluster of conspicuous long bristles (fig. 161). There is also a projection below the anal tube (formed by the conjunction of a process from each side) reaching beyond the latter {fig. 161) .............. trinervis (Beeker) Cheshire, Hereford., Lancs., Suffolk, N. Yorks. i,xi.

Process of left side of epandrium larger and with shorter hairs (figs. 164-165). There is no projection below anal tube .................. ~ .............................. 23

23 Hind metatarsus drawn out into apical, ventral, lobe so the spur lies below segment 2 of tarsus (fig. 162). Process ofteft side of epandrium larger, with curved posterior margin, and almost

34

subequal in length to process of right side (fig. 164) ................. collini Schmitz Cambs., Essex, Surrey, Sussex, Scotland: Roxburgh. iii-iv.

Hind metatarsus without apical, ventral,lobe (fig. 163). Process of left side of epandrium smaller, somewhat square-ended, and distinctly shorter than process of right side (fig. 165) •.......•................................................. papillats (Wingate) Widespread in British Isles. iii-ix.

Females (For distribution and phenology see males)

Many species possess a distinct stemite on abdominal segment 7. The form of this sternite is of great taxonomic value. It is advisable, therefore, to detach the abdomen, as close to the thorax as possible, and to mount it ventral side up beneath a separate coverslip (if necessary applying gentle pressure to the latter to ensure sternite 7 is wen exposed to view).

Vein 3 unforked, or with fork incomplete. In the latter case at least the basal part of vein 2 is absent, so that it is not attached to, or originating from, vein 3 (e.g; fig. 134) ... 2

Vein 3, forked (i.e. vein 2 always present and arising from vein 3) .................. 7 2 At least the distal third of vein 2 present. ....................................... 3

Vein 2 completely absent. .................................................... 5 3 Proboscis elongated, clearly reaching well beyond end of terminal bristles of palps. Legs largely

yellowish. Halteres yellow. V cin 3 about as wide as costal section 2 ............... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . minuta (Fabricius)

Proboscis not elongated. Legs mainly brown. Halteres brown to dark brown. Vein 3 with maximum breadth clearly greater than costal section 2 .......................... 4

4 Fore coxae largely yellowish, contrasting with brown femora. Wing membrane greyish with brownish (or darker greyish) tint in places, particularly beyond end of costa and bordering veins 4-6. The latter veins brown and readily discerned at low magnifications (e.g. x 10). Hairs of venter of segment 5 of abdomen strongly reduced in size, in marked contrast to normal-sized hairs on segments 4 and 6. At low magnifications segment 5 appears to be devoid of hairs altogether. ..................................... gracilis (Wood)

Fore coxae largely brown and not contrasting in colour with femora. Wing membrane uniformly greyish and veins 4-6 pale. Hairs on venter of segment 5 as strongly developed as those on segments 4 and 6 ................................ crassinervis (Strobl)

5 Fore coxae largely yellowish, contrasting with brown femora. Wing membrane greyish with brownish (or darker greyish) tint in places, particularly beyond end of costa and bordering veins 4-6. The latter veins brown and readily discerned at low magnifications (e.g. x 10). (See also couplet 4) ............................................ gracilis (Wood)

Fore coxae largely brown and not contrasting in colour with femora. Wing membrane uniformly pale greyish with veins 4-6 pale ..................................... 6

6 Hairs of abdominal venter at least as robust as those on posterior margin of tergite 6. With similar hairs on pleural region (behind spiracle) on segments 4-6, and usually with an isolated hair behind the spiracle on segments 2-3 as well ............... citreiformis (Becker)

Hairs of abdominal venter at most as robust as those on posterior margin of tergite 6. Pleural region devoid of hairs on segments 1-5 ......•..•..•.. .....•........ smithi Disney

7 Hairs of arista (in distal half) clearly shorter than maximum width of base of segment 3 of arista .............•......... , ................. " .......................... 8

Hairs of arista (in distal half) clearly longer than maximum width of base of segment 3 of arista .................................................................... 9

8 Hair at base of vein 3 approximately same length as costal cilia. Sternite of abdominal segment 7 as fig. 166. Halteres, legs and palps all dark brown to blackish ... Iugubris (Meigen)

Hair at base of vein 3 clearly longer than costal cilia. Palps yellowish brown (Note: I have not examined a female. The sternite of segment 7 has not been described) .......... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . subcompleta Schmitz

9 Halteres with at least knob yellowish to pale yellow ..........•................... 10 Halteres largely brown to blackish ............................................ 15

10 Vein 3 with unusually large fork and with 1-5 hairs on dorsal face before fork, these hairs being shorter and finer than bristle like hair at base of vein. Costa starts to broaden before midpoint of costal section 1 (fig. 148). (Posterior margin of venter of abdominal segment 6 with strongly developed, bristle-like hairs. Sternite of abdominal segment 7 as fig. 171.) ..................................... ; .........•......... autumnalis (Becker)

Vein 3 without fme hairs before fork. The latter usually clearly smaller, but if nearly as large then the costa starts to broaden beyond mid-point of costal section 1 ............. 11

35

11 Legs dominantly yellowish to yellowish-brown (Stemite of abdominal segment 7 as fig. 167.) ....................................................... Iuteifemorata (Wood)

Legs dominantly dark brown ................................................. 12 12 Stemite of abdominal segment 7 not developed. Abdominal venter conspicuously haired.

Fork of vein 3 small (as fig. 145) ............................. uudipalpis (Beeker) Sternite of abdominal segment 7 clearly developed, at least anteriody and posteriorly. Fork

of vein 3 usually longer .................................................. , .. 13 13 Stemite of abdominal segment 7 with two short posterior processes, one on each side (fig.

168) ........................................................ excisa (Lundbeck) Sternite of abdominal segment 7 with a single, median, posterior process (figs 169-170)

........................................................................ 14 14 Posterior process of stemite of abdominal segment 7 narrower and less extensively darkened

at tip (fig. 169) ............................................. ltyalinata (Meigen) Posterior process of sternite of abdominal segment 7 broader and more extensively darkened

at tip (fig. 170) ........................................... intempesta (Schmitz) 15 Scutellum with a posterior pair of bristles and an anterior pair of hairs which are clearly shorter

than hairs on scutum .......................•.............................. 16 Scutellum with 2 pairs of bristles. The anterior pair may be shorter than the posterior pair

but are always stronger than hairs on scutum ................................. 21 16 Axillary ridge of wing with 2 or more bristles. Sternite of abdominal segment 7 highly

distinctive (fig. 172). (Legs dominantly brown in colour) ......... flexipalpis Schmitz Axillary ridge of wing with a single bristle. Stemite of abdominal segment 7 not as in fig. 172

........................................................................ 17 17 Sternite of abdominal segment 7 ending in a median posterior process (fig. 167). (Legs

dominantly yellow to yellowish.) .......... '" .............. luteifemorata (Wood) Stemite of abdominal segment 7 of different shape or reduced .................... 18

18 Vein 3 at most as wide as distal half of costa (Sternite of abdominal segment 7 broadens posteriorly to a wide, more-or-Iess straight, hind margin) ............. vitrea (Wood)

Vein 3 clearly considerably wider than maximum breadth of costa ................. 19 19 Hairs of venter of segment 5 of abdomen strongly reduced in size, in marked contrast to

normal-sized hairs on segments 4 and 6. At low magnifications segment 5 appears to be devoid of hairs altogether. (see also couplet 4) ..................... gracilis (Wood)

Hairs of venter of segment 5 of abdomen as well developed as those on segments 4 and 6 ............................................•........................... 20

20 The posterior row of hairs on venter of segment 5 of abdomen little, if any, wider than rest of hair patch on this segment. ............................... antricola (Schmitz)

The posterior row of hairs on venter of segment S of abdomen about 3 x the width of rest of hair patch on this segment. The lateral extensions of this row of hairs reach well up the sides of the segment. ..................................... distinguenda (Strobl)

21 Stemite of abdominal segment 7 with 2 posteriorly directed processes (fig. 174). Hind tibia usually without a bristle in ~r two thirds ....................... opaca (Meigen)

Stemite of abdominal segment 7 not as fig. 174. Hind tibia usually with at least a small bristle on anterior face in upper two-thirds ....•.•.................................. 22

22 Axillary ridge of wing with only 1 bristle. Stemite of abdominal segment 7 not developed. Venter of abdomen densely clothed with strong, bristle-like, hairs .. triuervis (Becker)

Axillary ridge of wing with 2 or more bristles. Sternite of abdominal segment 7 present, but if somewhat inconspicuous the hairing of the venter is less robust and more-or-Iess restricted to a ventral median band (except on segment 6) •........••.................... 23

23 Anteriorly abdominal tergites 2-6 are differentiated into fields of discrete denticIes - fIg. 177 (seen in profile the posterior ends of these denticles project dorsally with a posterior inclination) ........•..................................................... 24

Abdominal tergites 2-6 normal (fig. 178). (Sternite of abdominal segment 7 a narrow lance­head shape (fig. 173). Proboscis with very robust labrum. Labella with conspicuous teeth. Axillary ridge of wing with less than 5 bristles.) ....•............... collini Schmitz

24 Stemite of abdominal segment 7 a broad lance·head shape with a blunt, black, upturned tip (fig. 175). Proboscis not abnormally developed, the labrum ending short of the tips of the pall's. Axillary ridge of wing with less than 5 bristles •.......... papillata (Wingate)

Stemite of abdominal segment 7 triangular with a broad, straiaht, posterior margin (fIg. 176). Proboscis strongly developed, with robust labrum which reaches beyond the tips of the

36

palps. Labella with teeth. Axillary ridge of wing more than 3 bristles (usually) 7 .... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . intermedia (Malloch)

Genus Woodiphora Schmitz (Figs. 179-180)

This genus includes more than 20 species from all regions of the world. 3 species are known from the Palaearctic Region.

The larval natural history is not known. The data label attached to a specimen in the Verrall-Collin collection implies it was associated with the excavation into a tree produced by the caterpillar of the Carpenter Moth (Cossus cossus)

A dark species. Halteres dark. Mesopleuron bare. Scutellum with 4 strong bristles. Vein 2 more-or-Iess perpendicular to costa (fig. 179). C1 , with large palps bearing short hairs, and only short bristles at their tips. Hypopygium as fig. 180 .......... retroversa (Wood) Essex, Kent, Hants., Hereford. vii-viii.

References

BORGMEIER, T. 1963. Revision of the North American Phorid Flies. Part 1. (Dipt. Phoridae). Studia Ent., Petropolis 6: 1-256.

CARTER, J .B. 1977. Survival of Tipu/a paludosa (Dipt., Tipulidae) larvae after infection with larvae of Megase/iapaludosa (Dipt., Phoridae). Entomologist's mono Mag. 112: 245 (1976)

COGGINS, R.E. 1970. Megaselia paludosa (Wood) (Dipt., Phoridae) parasitising Tipu/a paludosa Mg. (Dipt., Tipulidae). Entomologist's mono .Mag. 106: 108.

COL YER, C.N. 1954a. The "Coffin" fly, Conicera tibialis Schmitz (Dipt., Phoridae). J. Soc. Brit. Ent. 4: 203-206.

COLYER. C.N. 1954b. More about the "Coffin" fly. Conicera tibialis Schmitz (Diptera, Phoridae). Entomologist 87: 129·132.

COL YER, C.N. 1954c. Further emergences of Conicera tibialis, the 'coffin' fly (Diptera. Phoridae). Entomologist 87: 234.

COL YER, C.N. 1954d. 'Swarming" of Phoridae (Diptera). J. Soc. Brit. Ent. 5: 22·27 DECOU-BURGHELE, A. 1961. Sur la biologie de Megaselia melanocephala von Roser, phoride

parasite des cocons de Meta menardi latr. Annal. Lab. souterrain Han·sur·Lesse (Belgium) 11: 16-22.

DELAGE, A. & LAURAIRE, M·CL. 1974. Mise au Point sur le genre Phalacrotophora (Dipt., Phoridae) et description de I'ovipositor d'une femelle parasite. Annals parasi!. hum. comp. (Paris) 49: 495·500.

DISNEY, R.H.L. 1975. A key to the larvae, pupae and adults of the British Dixidae (Diptera) the meniscus midges. Freshwat. Bioi. Assoc. Sci. Publ. 31: 1·78.

DISNEY. R.H.L. 1976. A further case of a nematoceran fly (Diptera: Sciaridae) parasitised by a species of scuttle fly (Diptera: Phoridae). Entomologist's Gaz. 27: 91·98.

DISNEY, R.H.L. 1977. A further case of a scuttle fly (Dipt. Phoridae) whose larvae attack slug eggs. Entomologist's mono Mag. 112: 174 (1976).

DISNEY, R.H.L. 1978. A new species of afrotropical Megase/ia (Diptera: Phoridae), with a re­evaluation of the genus Plastophora. Z. ang. Zool. 65: 313-319.

DISNEY, R.H.L. 1979a. Spiniphora helicivora (Dufour) is a synonym of S. maculata (Meigen) (Dipt., Phoridae). Entomologist's mono Mag. 113: 201-202 (1977).

DISNEY, R.H.L. 1979b. Natural history notes on some British Phoridae (Diptera) with comments on a changing picture. Entomologist's Gaz. 30: 141-150.

DISNEY, R.H.L. 1979c. The British Metopina (Diptera: Phoridae) with description of a new species. Zool. J. Linnean Soc. 67: 97·113.

DISNEY, R.H.L. 1979d. A new species of Phalacrotophora (Dipt., Phoridae) from France. Annals Parasit. hum. comp. (Paris) 54: 533·536.

DISNEY, R.H.L. 1980a. Records of flower·visiting by scuttle flies (Diptera: Phoridae) in the British Isles. Naturalist, Hull 105: 45·50.

37

DISNEY, R.H.L. 1980b. Variation in Megaselia pulicaria (Fall.) (Dipt., Phoridae) with the recognition of new synonymies. Entomologist's mono Mag. 115: 97-103 (1979).

DISNEY. R.H.L. 198Oc. A scuttlefly (Dipt., Phoridae) that parasitises larvae of Trichosia Winnertz (Dipt., Sciaridae). Entomologist's mono Mag. 116: 5-6.

DISNEY, R.H.L. 1980d. A new species of Gymnophora Macquart (Diptera Phoridae) from Yorkshire and Durham. Naturalist, HulllOS: 125-131.

DISNEY, R.H.L. 1980e. Chaetopleurophora bohemanni (Seeker) (Diptera: Phoridae) added to the British List. Efltomologist's Gaz. 31: 245.

DISNEY, R.H.L. 1981a. A further synonym in the genus Conicera Meigen with a revised list of the British species. (Diptera: Phoridae). Entomologist's Rec. J. Var. 93: 126-128.

DISNEY. R.H.L. 1981 b. An exotic scuttlefly Chonocephalus heymonsi Stobbe (Dipt., Phoridae) from Middlesex. Entomologist's mono Mag. 116: 207-212. (1980).

DISNEY, R.H.L. 1981e. A fourth species of Gymnophora (Dipt., Phoridae) from England. Entomologist's mono Mag. 116: 214 (1980).

DISNEY, R.H.L. 1981d. A new, and first Afrotropica1, species of Auxanommatidia (Diptera: Phoridae) Z. ang. Zool. 67: 323-330 (1980).

DISNEY, R.H.L. 1981 e. A sixth species of Borophaga Enderlein (Dipt., Phoridae) for the British List. Entomologist's mono Mag. 117: 141.

DISNEY, R.H.L. 1981f. What is Diplonevra versicolor (Sehmitz) (Dipt., Phoridae)? Entomologist's mono Mag. 117: 157-158.

DISNEY, R.H.L. 1982a. A seuttlefly (Dipt., Phoridae) new to Britain Entomologist's mono Mag. 117: 178 (1981).

DISNEY, R.H.L. 1982b. Theunderscribed female and first British record of Metopina crassinervis Sehmitz (Dipt., Phoridae). Entomologist's mono Mag. 117: 179-182 (1981).

DISNEY, R.H.L. 1982c. A scuttle fly (Diptera: Phoridae) that appears to be a parasitoid of a snail (Stylommatophora: Zonitidae) and is itself parasitised by a Braconid (Hymenoptera). Entomolgist's Ree. J. Var. 94: 151-154.

DISNEY, R.H.L. 1982d. What is Diplonevra abdominalis (Fallen) (Dipt., Phoridae)? Entomologist's mono Mag. 118: 113-115.

DISNEY, R.H.L. 1982e. Confirmation of a suspected synonym in the genus Borophaga (Diptera: Phoridae) Entomologist's mono Mag. 118: 115.

DISNEY, R.H.L. 1982f. A new species of Phora (Dipt., Phoridae) from Ireland. Ir. Nat. J. 20: 425-426.

DISNEY, R.H.L. 1982g. Re-eva!uation of the genus Citrago Schrnitz (Dipt., Phoridae) and a consequent secondary homonym. Entomologist's mono Mag. 118: 247-248.

DISNEY, R.H.L. 1982h. The true identity of the Triphleba polposa (Dipt., Phoridae) ofthe British List. Entomologist's mono Mag. 118: 255-256.

DISNEY, R.H.L. 1982i. A species of Phora (Diptera: Phoridae) new to the British List. Entomologist's Gaz 33: 257.

DISNEY, R.H.L. 1983. Triphleba subeompleta Schrnitz (Dipt., Phoridae) new to Britain. Entomologist's mono Mag. (in press).

DISNEY, R.H.L. COULSON, J.C. & BUTTERFIELD, J. 1981. A survey of the scuttIeflies (Diptera: Phoridae) of upland habitats in Northern England. Naturalist, Hull 106: 53-66.

DISNEY, R.H.L. & EVANS, R.E. 1980. Phoridae (Diptera) whose larvae feed on eggs of spiders (Araneida). Entomologist's mono Mag. 115: 21-22 (1979).

DISNEY, R.H.L., HENDERSON, I.F., PERRY, J.N. & CLEMENTS, R;O. 1981. Phoridae (Diptera) from English Pasture soils. Pedobiologia 22: 366-378.

DISNEY, R.H.L. & SMITH, K.G. V. 1978. Scuttle flies (Diptera, Phoridae) that breed in man. Roy. Soc. trop. Med. Hyg., Symposium Proe. Med. Ent. Centenary, 1977. (Entomol. Demonstrations) p. 136.

DONISTHORPE, H. St. J .K. 1927. The Guests of British Ants - their Habits and life-histories. Routledge, London.

FULLER, M.E. 1934. The early stages of Sciadoeera rufomaculata White (Dipt., Phoridae). Proc. Linn. Soc. N.S. Wales 59: 9-15.

GOETSCH, W. 1940 Vergleiehende Biologie der Insekten Staden. Leipzig. GRIFFITHS, G.C.D. 1972~ The Phylogenetie ClassifICation ofDiptera Cye/orrhapha. Junk, The

Hague. HARDY, A. 1965. The Living Stream. Coilins, London. HENNIG, W. 1971. Zur Situation der biologsichen Sytematik. Erlanger Forseh. (8) 4: 7-15.

38

HENSHA W, D.l. de C. 1981. Observations on the preparation of Berlese's Fluid. Entomologist's man. Mag. 116: 206 (1980).

KLOET, G.S. &HINCKS, W.D. 1976. A check list of British Insects. (Second edition). Diptera and Siphonaptera. Handbk. Ident. Br. Insects 11(5): 1-139.

KLOTER, K.O. PENNER, L.R. & WIDMER, W.l. 1977. Interactions between the larvae of Psychoda alternata (Dipt., Psychodidae) and Dohrniphora cornuta (Dipt., Phoridae) in a trickling filter sewage bed, with descriptions of the immature stages of the later. Ann. Entomol. Soc. Am. 70: 775-781.

LEWIS, D.J. 1973. in SMITH, K.G.V. (Bd). Insects and other Arthropor]sofMedicallmportance. British Museum (Natural History), London.

LUNDBECK, W. 1922. Diptera Danica Part VI Pipunculidae, Phoridae. Wesley, London. McALPlNE, 1.F. & MARTIN, 1.E.H. 1966. Systematics of Sciadoceridae and relatives with

descriptions of two new genera and species from Canadian amber and erection of family Ironomyiidae (Diptera: Phoroidea). Canad. Ent. 98: 527-544.

McALPlNE, J.F., PETERSONB.V., SHEWELL, G.E., TESKEY, H.l., VOCKEROTH, J.R., & WOOD, D.M. 1981. Manual of Nearctic Diptera Volume 1. Agriculture Canada, Ottawa.

McCRAE, A.W.R. 1967. Infestation of emulsion paint by the fly Megase/ia scalaris (Loew) (Dipt., Phoridae). Entomologist's mono Mag. 102: 241-243.

MORRIS, H.M. 1922. On the larva and pupa of a parasitic phorid fly, Hypocera incrassata Mg. Parasitology 14: 70-74.

OLDROYD, H. 1964. The Natural History of Flies. Wiedenfeld and Nicolson, London. OLDROYD, H. 1970. Diptera 1. Introduction and Key to families. Handbk. Ident. Br. Insects

9(1): 1-104. PALKA, J., SCHUBIGER, M. & HART, H.S. 1981. The path ofaxons in Drosophila wings in

relation to compartment boundaries. Nature Lond. 294: 447-449. PICARD, F. 1930. Sur le parasitisme d'un Phoride (Megaselia cuspidata Schmitz) aux depens

d'un myriapode. Bull. Soc. zool. Fr. SS: 180-183. ROBINSON, W.H. 1971. Old and new biologies of Megaselia species (Diptera, Phoridae). Studia

Ent., Petropolis. 14: 321-348. ROBINSON, W.H. & FOOTE, B.A. 1968. Biology and immature stages of Megaselia aequalis,

a phorid predator of slug eggs. Ann. ent. Soc. Americ. 61: 1587-1594. SCHMITZ, H. 1929. Diptera of Patagonia and South Chile Part VI, Fascicle 1- Sciadoceridae

and Phoridae. British Museum (Natural History), London. SCHMITZ, H. 1938a. Die ersten hundert Phoriden von Portugal. Broteria 34: 163-179. SCHMITZ, H. 1938b. - 1958. 33. Phoridae. In LINDNER, E. (Bd). Die Fliegen der

Palaearktischen Region. Schweizerbart'sche, Stuttgart. SCHMITZ, H. & BEYER, E. 1965 - 1974. Ibid. continued. SCHMITZ, H. & DELAOE, A. 1974 - 1981. Ibid. continued. SCHMUTTERER, H. 1952. Plastophora rufa (Wood) (Dipt., Phoridae) als Eirauber and Parasit

von Eulecamium corni (Bche) (Homopt., Coccoidea). Auz. Schadlingsk.15: 145-14J!. SMITH, K.O.V. 1977. Notes on some British Phoridae (Diptera) including two species of Megaselia

Rondani new to science. Entomologist's Rec. J. Var. 89: 161·168. SOUTHWOOD, T.R.E. 1980. Temporary mounts for specimens in alcohol. Antenna 4: 2. TUXEN, S.L. 1969. Nomenclature and homology of genitalia in insects. Memorie Soc. ent. ital.

48: 6-18. VAN EMDEN, F.!. 1950. Dipterous parasites of Coleoptera. Entomologist's mono Mag. 86:

182-206. VENTURI, F. 1966. La struttura morfologica Dell'Ipopigio di un dittero Foride (Triphleba

antricola Schrnitz). Frustula Entomoigica. 9: 1-42. WESCHE, W. 1908. The systemantic affinities of the Phoridae and of several Brachycerous families

of Diptera. Trans. ent. Soc. Lond. 1908: 283-296. Y ARKULOV. F. 1972. Mukha - gorbatka Phora holosericea (Diptera. Phoridae) - khishchnik

kornevykh tiey. Zool. Zh. (U.S.S.R.) 51: 1415-1418.

39

Plates and figures

The figures have all been prepared from slide-mounts of specimens. In every case the scale lines = 0.1 mm.

The following figures have been published previously in the journals indicated:-

Figs. 8,25, 28-30, 57, 76-78, 92, 133 Figs. 62-63, 65-67, 69, 71-72 Figs. 99-100

Figs. 106, 126

in the Entomologist's Monthly Magazine. in the Naturalist. in the Annales de Parasitologie (Masson S.A., Paris) in the Irish Naturalists' Journal.

CA UTION: In matching specimens against drawings of complex three-dimensional structures, such as the male hypopygium, allowance must be made for alterations in appearance resulting from slight shifts in orientation. Also the hypandrium may be more or less displaced depending on the degree of protrution, or retraction, of the penis complex.

40

The microsculpture of the posterior face of middle of hind femur in Diplonevra nitidula 9 -Plate A, and D. glabra 9 - Plate B. (For scale see fig. 60).

41

1

2

-..: ~ .... '

.- ...... !-r ..

-,-. - ~ P:" ."_._ ... _~.

_. ~~o..'~"::"~ .. :~

Figs 1-5. Pre-adult stages. 1. egg of Megaselia ha/terata. 2. egg of Megaselia ruflCornis in side view. 3. larva of Dohrniphora comuta. 4, larva of Megaselia rujipes. 5. puparium of M. rujipes (at start of adult eclosion).

42

cercus~

anal / tube

proctiger

1

co

BB Se I

h/""'----"

8

FIgS 6-8. Adult morphology. 6, head of Triphleba nudipalpis - 0 = ocellar bristles, PL ,.. postero­laterals, PO = pre-ocellars, ML ,.. medio-laterals, AL = Antero-Iaterals, A ,.. antials, SA = supra-antennals. 7, tenninalia of male abdomen (end of segment 6 plus hypopygium) of Megaselia type (generalised) viewed from left side- T. 6 = Tergite 6.8, wing of Diplonevraflorea. t-7 ,.. longitudinal veins, CO - Ct = section 1 of costa, Cl - C2 ,.. section 2, C2 - C3 = section 3, CC = costal cilia, BB ,.. basal bristle of costa, AB ,.. bristles of axillary ridge, h = humeral vein, SC ,.. subcostal vein.

43

\-' St

1---1 ...,..-:~ ::y, ... . r? c ,~c . .

Hy

10

Figs 9-11. Adult morphology continued. 9, thorax of Megaselia ciliata viewed from left side - hu = humerus, s = spiracle, Pr = propleuron, Me = mesopleuron (f = mesopleural furrow), Pt = pteropleuron, St = stemopleuron, Hy = hypopleuron, ha = haltere, c = coxa. 10, thorax of Conicera dauci labelled as fig. 9. 11, foretibia and tarsus of Spiniphora bergenstammi -s = spur. 12, hind tibia of Pseudacteon brevicauda - P = hair palisade.

44

,:!.~;;,J.;'!,; r~J: ~!:;:-':~ '(,;, ;':,\,':;~.:>:';

e,,;,j\y':'~';::<:::' f;\:;;,:J\·~>,'~~.:::;:~

~ (::) :~.'; ':i.:~ :';:~';":J:::':': ,~,,', :~.,:,:':",',::;\ .'~:::

15

Figs 13-17. Aenigmatiasspp. 13, wing of A.Jranzi Cl. 14-17 A.lubbQcki 14, wing of Cl. 15, female (legs omitted). 16, hypopygium from left side. 17, hypopgium from right side.

4S

- ..... ~.

~. -~ .....

--.' . . ----------. .".

21 ~ ..• ..w ..• ~.~.<~:.""~'.:.' .... ~ .... :.~. J .. . : ...... ' .' ' .. :,'>~

• ," co .• .. ' . '." .:.' .

. .,- I •

,:::., . ~ ". ,. ... ;', .:'- ' "

..... :: ..... ~ ... ~ .

-~~ ..• - ... -.: .. . . ....

. -:... ~

2Z'~~_~_ Figs 18-22. 18, Anevrina thoracica Cl anterior face of base of hind femur. 19, Anevrina curvinervis

Cl Posterior face of base of hind femur. 20, Beckerina umbrimargo Cl wing. 21, Borophaga irreguiaris 0' distal extremity of costa and vein 3. 22, Borophaga subsultans Cl distal extremity of costa and vein 3.

46

<€l t .~

.~ it (if I,' ,.

I:' .:! (@ " ~.

23

Figs 23-27.23. Borophaga incrassata 0' frons (bristles omitted). 24. Borophaga carinifrons 0'

frons (bristles omitted). 25. Chonocephalus heymonsi 0' wing. 26-27 Plectanocnema nudipes 0'. 26, anterior face of hind tibia. 27, hypopygium from left side.

47

I

Figs 28-30. Chonocephaius heymonsi. 28, 0' hypopygium from right side. 29, female (legs omitted). 30, 9 hind leg.

48

I

I I I

I I 36

Figs 31·39. Conicera spp. er er 31, C. dauciIeft clasper of hypopygium from side. 32-37 right c1asper of hypopygium, external face. 32, C. dauci. 33, C. tibialis. 34, C. simils. 35, C. tarsa/is. 36, C. schnittmanni. 37, C. j7oricola. 38·39 Posterior face of mid femur. 38, C. tibialis. 39, C. similis.

49

I I I 40

I

Figs 40·45.40·43 Conicera spp. Q Q, anterior face of hind tibia (hairs omitted). 40, atypical C. similis. 41, C. dauci. 42, C. tibialis. 43, C. jloricola. 44-45 Oymnoptera spp. Cl 0', posterior face of base of hind femur. 44, O. /ongicostalis. 45, O. vitripennis.

50

Figs 46-50. Dip/onevra spp. 0' 0' Posterior face of base of hind femur and trochanter. 46, D. abbreviata. 47-48, D. f/orea. 49, D. nitidu/a. SO, D. junebris.

51

;"

53 11-----

Figs 51·55. Diplonevra spp. et et 51-53, posterior face of base of hind femur and trochanter. 51, D. concinna. 52, D. glabra. 53, D. pilose/la. 54-55, hypopygium from right side. 54, D. glabra. 55, D. pilosella.

52

4

-'

58

59

51

.5

--- -.r---;

s· :-­~

Figs 56-60. Dipionevra spp. 56, D. abbreviate 9 wing. 57, D. florea 0' wing. 58, D. funebris 9 segments 4-6 of abdomen in side view - s = spiracle. 59, D. pi/oseJJa 9 tip of right labellum of proboscis from above. 60, D. nitiduia 9 microsculpture of posterior face of hind femur.

53

Figs 61-62 Gymnophora spp. Cl hypopygium from left side. 61, G. arcuata. 62, G. quartomollis.

54

~--

-~i ~r~~

-

63

Figs 63-64. Oymnophora spp. Cl bypopygium from left side. 63, O. hea/eyae. 64, O. integra/is.

55

I

66

&9~ ----

Figs 65-69. Gymnophora spp. 65, G. healeyae 0' top half of thorax from left side - H = humerus, s = spiracie, M = mesopleruron, N = notopleural bristles, W = wing base. 66-68, « abdominal tergite 8.66, G. quartomollis. 67. G. healeyae. 68, G. integralis. 69, G. healeyae « wing.

56

73

71

8.··· .. · ... · .. · .... · .... ·

U 1

72

\ \ \

\ ,'/

I

Figs 70-75.70-74 Gymnophora spp. 9 9 abdominal tergites and associated internal sacs. 70, G. quartomollistergites 5 and 6.71, G. healeyaetergite 6.72, G. healeyaetergite 7. 73, G. integralis. tergites 5 and 6. 74, G. integralis tergite 7. 75, Hypocera mordellaria 9 anterior face of hind tibia.

57

79 80

Figs 76-82. Metopina spp. 76-77 M. crassinvervis. 76, Q wing. 77, er wing 78, M. heselhausi er wing. 79-82 hair patch on segment 4 of abdominal venter of er. 79, M. heselhausi. 80, M. oligoneura. 81, M. pileata. 82, M. perpusilla.

58

83

Figs 83-88. Metopina spp. 0' 0'. 83, M. ulrichi posterior face of hind femur. 84-88 posterior face of base of hind femur and trochanter 84, M. heselhausi. 85, M. oligoneura. 86, M. pi/eata. 87, M. braueri. 88, M perpusilla.

59

......... ~ ..... ~.;~:.:;.;.:~~ .......... ~.

I

. ... -......... ~' ... ~.

I ~.: ... -•.. ~\ ..• '- .. '-j \' r

1 r

93

., ,

It! f!;/

96

Figs 89·96. Metopi'na spp. 9 9. Abdominal tergites 4 and 5.89, M. galeata. 90, M. braueri. 91, M. ulrichi. 92, M. crassinervis. 93, M. perpusilla. 94, M. pileata 95, M. heselhausi. 96, M. oligoneura.

60

99 >----<

---.::J 101 .-4

Figs 97·100. Phalacrotophora spp. 97·98, er hypopygium from left side. 97, P. berolinensis. 98, P. fasciata. 99·100, <;> ovipositor from left side. 99, P. berolinensis. 100, P. fasciala.

61

101

t-----I

183

184

Figs 101-107.101, Dohrniphoracornuta CJ' posterior face of base of hind femur. 102-107 Phora spp. CJ' hypopygium viewed from left side (on left) and from right side (on right). 102, 103, P. dubia. 104, 105, P. artifrons. 106, 107, P. speighti.

62

108

112

114

Figs 108-115. Phora spp. et hypopygium viewed from left side (on left) and from right side (on right). 108, 109, P. atra. 110, Ill, P. holosericea. 112,113, P. edentata. 114, 115, P. tincta.

63

I 118

122

Figs 116-123. Phora. spp. et hypopygium viewed from left side (on left) and right side (on right). 116,117, P. stictica. 118,119, P. praepandens. 120, 121, P. hamata. 122, 123, P. obscura.

64

124

127

129

4lJ ,~

iJ ,()

'. • " 0 • . .,

, . .: ... '~~'.----. :;' ~.

125

.~.,,". .

tJ'

126

128 /

Figs. 124-U9. 124-126 Phora spp. 0' frons. 124, P. artifrons. 125, P. praepanliens. 126. P. speighti. 127"129 Pu/iciphora borinquenensis .. 127, 0' wing. 128, <:;( abdominal tergites 4-6. 129, 0' hypopygium viewed from left side.

65

I

130

13l

I 133

134 ..-

Figs 130-134. 130-133. Spiniphora spp. 130, 131 S. excisa.130, right palp and antenna of et . 131, palp of <;.>. 132, 133, S. bergenstammi. 132, right palp and antenna of 133, palp of <;.>.134, Triphleba crassinervis et wing.

66

131

138

.~ .....

1~~0"C<' Figs 135-139. Triphleba spp. 135-138 right antenna of 0". 135, T. citreijormis. 136, T. minuta.

137, T.smithi. 138, T. crassinervis. 139, posterior face of base of hind femur of T. excisa. 0".

67

141

141

142

Figs 140-144. TriphJeba spp. Cl hypopygia. 140-142, process of upper half ofleft side of epandrium. 140, T. citreiformis. 141, T. smithi. 142, T. crassinervis. 143-144, entire hypopygium of T. crassinervis. 143, left side. 144, right side.

68

145

...• ~ .... 147 ~ ~.., '.

.... ~

··~.··C:':::z;2~ .... ~ ......... ::~ ..... ~

148'" .....• .'. . .... ~. ". ".

~ ~. ",.' .' .~ . ... :~., '.

149 .... ..... ~ ".

Figs 145·149. Triphleba spp. anterior, basal part of right wing (costa and thick veins only). 145, T. nudipalpis Cl. 146, T. hyalinata Cl. 147, T. autumnalis Cl. 148, T. autumnalis 9. 149, T. intempesta Cl.

69

150

152

Figs 150-152. Triphleba spp. CY' hypopygia. 150, T. luteifemorata left side. 151-152, T. hyalinata. 151, left side. 152, right side.

70

155

157

158

Figs 153-158. Triphleba. spp. et et. 153-154 T. intempestahypopygium. 153, left side. 154, right side. 155-157 T. f/exipalpis. 155, left side of hypopygium with penis complex fully extruded. 156, palp, with outline of antenna drawn to same scale. 157, right side of hypopygium (with penis complex omitted). 158, T. vitrea left side of epandrium.

71

159

160

Figs 15!J.161. Triphleba spp. Cl' bypopygia. 159, T. opaca terminal part ofleft side of epandrium. 160. T. intermedia terminal part of left side of epandrium. 161, T. trinervis left side.

72

162

163

1&4

185

Figs 162·165. Triphleba spp. 0' 0' •. 162·163 anterior face ofi\ind metatarsus. 162, T. collini. 163, T.papillata. 164-165terminalpartofJeftsideofepandriUll1.1~, T. collini. 165, T.papillata.

73

166 187 168

1

169 170

Figs 166-170. Triphleba spp. <;> abdominal sternite 7. 166, T. lugubris. 167, T. IUleijemorata. 168, T. excisa. 169, T. hyalinata, 170, T. inIempesta.

74

1

111 112

113

114 175 116

f'igs rn,·176" Triphlebu ,pp. 9 abdominalsternite 7. 171, T. autumnalis 172, T.jlexipalpis, 173, T. co!lim, \74, T opaca. 175, T. papillata. J 76, T, inrermedia,

75

117

118

179

180

Figs 177-180.177-178 Triphleba spp. 9 abdominal tergite 4. 177, T. papillata. 178, T. collini. 179-180 Woodiphora retroversa CT. 179, right wing.·180, left side of hypopygium.

76

182

Figs 181·183. r::t hypopygia. 181-182, Phora builata. 181. from left side. 182, from right side. 183, Triphleba lugubris from left side.

77

185

186

Figs 184-186. Triphleba spp. CY hypopygia. 184, T. /ugubris from right side. 185-186, T. subcomp/eta. 185, from left side. 186 from right side.

78

Index Principal references are in bold. Synonyms are in italics.

abbreviata (Diplonevra) 13,22,23 abdominalis (Diplonevra) 13 Aenigmatias 12, IS, 17 ajricana (Puliciphora) 14 agilis (Borophaga) 12, 20 albipennis (Conicera) 13 albispina (Triphleba) 14 albohalterata (Pseudacteon) 14 amplicornis (Diplonevra) 13 Aneurina 12 Anevrina 12, 15, 18 antricola (Triphleba) 14, 34. 36 aptina (Triphleba) 14 arcuata(Gymnophora) 13, 24 artifrons (phora) 14, 29 aterrima (Phora) 14 atra (Conicera) 13 atra (Diplonevra) 13 atra (Phora) 14, 29 atricapilla (Phalacrotophora) 14 autumnalis (Triphleba) 14, 33, 35

bartholomei (Triphleba) 14 basalis (Dohrniphora) 13 Beckerina 12, 17,19 bequaerti (Dohrniphora) 13 bergenstammi (Spiniphora) 14,32 bergenstammi (Spiniphora) 14 berolinensis (Phalacrotophora) 14, 28 blattoides (Aenigmatias) 12 bohemanni (Chaetopleurophora) 12, 20 borinquenensis (Puliciphora) 14, 31 Borophaga 12, 15, 19 braueri (Metopina) 13,27 brevicauda (Pseudacteon) 14,31 brevifrons (Aenigmatias) 12, 18 bullata (Phora) 14, 29

ca/iginosa (Anevrina) 12 carjnifrons (Borophaga) 12, ;W Chaewpleurophora 12, 15, !6, ;W chlorugastra (Dohrniphora) 13 Chonocephalus 12, 15, 17,10 ciliata (Megaselia) 44 cimbicis (Diplonevra) 13 Citrago 14 citreiformis (Triphleba) 14, 32, 35 cleghorni (Dohrniphora) 13 collini (Triphleba) 14, 34, 36 collini (Triphleba) 15 comstocki (Spiniphora) 14 concinna (Diplonevra) 13, 22, 23 concinna (Diplonevra) 13 conc%r (Triphleba) 14

Conicera 13, 17,21 connexa (Triphleba) 15 cornuta (Dohrniphora) 13, 24 crassicornis (Diplonevra) 13 crassinervis (Metopina) 13,26,27 crassinervis (Triphleba) 14, 33, 35 crockeri (Dohrniphora) 13 cuneata (Metopina) 13 curvinervis (Anevrina) 12, 18, 19

dauci (Conicera) 13, 21 debilis (Gymnophora) 13 dimidiata (Anevrina) 12 Diploneura 13 Diplonevra 13, 16,22 disparinervis (TriphJeba) 15 distincta (Diplonevra) 13 distinguenda (Triphleba) 14, 34, 36 divaricata (Dohrniphora) 13 Dohrniphora 13, 16,23 domestica (Spiniphora) 14 dorni (Aenigmatias) 12 dorsalis (Spiniphora) 14, 31, 32 dubia (ph ora) 14,29

edentata (Phora) 14,29 enervata (Triphleba) 14 erythronota (Chaetopleurophora) 12,20 excisa (Spiniphora) 14, 32 excisa (Triphleba) 14, 33, 36

jal/ens (Conicera) 13 jantinii (Triphleba) 14 fasciata (Phalacrotophora) 14,28 femorata (Borophaga) 12, 19 jemorata (Borophaga) 12 jennica (Anevrina) 12 jlavicornis (Spiniphora) 14 jlavimana (Borophaga) 12 jlavipaipis (Hypocera) 13 jlaviventris (Dohrniphora) 13 flexipalpis (Triphleba) 14, 34, 36 flexuosa (Diplonevra) 13 florea (Diplonevra) 13,22,23 floricola (Conicera) 13, 21, 22 jorcipata (Triphleba) 14 formicarum (Pseudacteon) 14, 31 franzi (Aenigmatias) 12, 18 jratercula (Triphleba) 15 fraudans (Dohrniphora) 24 julva (Dohrniphora) 13 julvipalpis (Conicera) 13 julvivemris (Diplonevra) 13 funebris (Diplonevra) 13, 22, 23

79

galeata (Metopina) 13, 26, 27 gaieata (Metopina) 13 genita/is (Gymnoptera) 13 gilsoni (Tripbleba) 15 glabra (Diplonevra) 13, 23 gracilis (Triphleba) 14,33,34,35,36 Gymnophora 13, 17, 24 gymnophorina (Diplonevra) 13 Gymnoptera 13, 17, 2S

halterata (Megaselia) 42 hamata (phora) 14,30 hamata (Phora) 14 healeyae (Gymnophora) 13,24,25 helicivora (Spiniphora) 14 heselhausi (Metopina) 13, 26, 28 heterocerca (Phora) 14 heymonsi (Chonocephalus) 12, 20 hiemalis (Triphleba) 14 highlandicus (Aenigmatias) 12 holosericea (Phora) 14, 29 hyalinata (Tripbleba) 14, 34, 36 hyemalis (Tripbleba) 14 Hypocera 13, 15,25

immaculata (Spiniphora) 14 inaequalis (Metopina) 13 incrassata (Borophaga) 12, 20 integralis (Gymnophora) 13,24,25 intempesta (Tripbleba) 14, 34, 36 intermedia (Tripbleba) 15,34,37 intricata (Spiniphora) 14 irregularis (Borophaga) 12, 19

jamaicensis (Chonocephalus) 12

lehmanni (Tripbleba) 15 longicostalis (Gymnoptera) 13,25 lubbocki (Aenigmatias) 12, 18 lubbocki (Aenigmatias) 12 luctuosa (Diplonevra) 13 lugubris (Triphleba) 15,33,35 lugubris (Triphleba) IS luteifemorata (Tripbleba) IS, 33, 34, 36

maculata (Spiniphora) 14,31, 32 maculata (Spiniphora) 14 Megaselia 3, 13, 16 melanocephala (MegaseIia) Metopina 13, 16,25 minor (pseudacteon) 14 minuscula (Conicera) 13 minuta (Tripbleba) IS, 32, 35 mohrae (Triphleba) 14 montana (Phora) 14 mordax (Dohrniphora) 13 mordellaria (Hypocera) 13, 25

navigans (Dohrniphora) 13

80

nevadae (Metopina) 13 nickerli (Conicera) 13 nigricornis (Tripbleba) 15 nigrocincta (Phalacrotophora) 14 nigrodorsata (Chaetopleurophora) 12 nitidijrons (Diplonevra) 13 nitidula (Diplonevra) 13,22,23 nitidula (Diplonevra) 13 notata (Spiniphora) 14 nudipalpis (Tripbleba) 15,33,36 nudipes (Plectanocnema) 14, 30 nuptamjerens (puliciphora) 14

obscura (Dohrniphora) 13 obscura (phora) 14,30 occidenta/is (puliciphora) 14 octobris (Triphleba) 14 o'kellyi (Borophaga) 12 oligoneura (Metopina) 13,26,27 opaca (Tripbleba) 15, 34, 36 opaca (Triphleba) 14, IS opposita (Dohrniphora) 13

pa/pina (Diplonevra) 13 papilIata (Triphleba) 15,34,36 parcepilosa (Diplonevra) 13 pauxilla (Conicera) 13 perennijormis (Triphleba) 14 perennis (Triphleba) 14 perpusilIa (Metopina) 13, 27 Phallacrotophora 14, 16,28 Phora 14,17,28 pileata (Metopina) 14, 26, 27 piloseIIa (Diplonevra) 13,23 Platyphora 12 Plectanocnema 14, 16,30 praepandens (phora) 14, 30 pressata (Spiniphora) 14 Pseudacteon 14, 16,30 pseudoconcinna (Diplonevra) 13 pubericomis (Triphleba) 15 Puliciphora 14, 15, 17, 31 punctijascia (Chonocephalus) 12

quadrata (Anevrina) 12 quartomollis (Gymnophora) 13, 24

retroversa (Woodiphora) 15,37 rhenana (Metopina) 14 rostralis (Diplonevra) 13 ruficornis (Megaselia) 42 rujicornis (Tripbleba) 15 rufipes (Gymnophora) 13 rufipes (Megaselia) 42 rufipes (puliciphora) 31

schineri (phora) 14 schnittmanni (Conicera) 13, 21, 22 similis (Conicera) 13. 21, 22

similis (Conicera) 13 similis (Triphleba) 15 smithi (Tripbleba) 15, 33, 35 sordidipennis (Diplonevra) 13 sororcula (Diplonevra) 13 speighti (phora) 14, 30 sphigicides (Spiniphora) 14 sphingicides (Spiniphora) 14 spinicollis (Puliciphora) 14 Spiniphora 14,17,31 spinosior (Chaetopleurophora) 12,20 spinosissima (Chaetopleurophora) 12,20 stictica (phora) 14, 29 stictica (Phora) 14 subcompleta (Triphleba) 15,33,35 sublugubris (Tripbleba) 15 sub sultans (Borophaga) 12, 19 subsultans (Hypocera) 13 suspecta (Tripbleba) 15

tarsalis (Conicera) 13,21, 22 termitophila (puliciphora) 14 thoracica (Anevrina) 12, 18 thoracica (Spiniphora) 14 tibialis (Conicera) 13, 21, 22 tincta (Phora) 14,30

trinervis (Triphleba) 15, 34, 36 Tripbleba 14, 17, 32 trochanterata (Anevrina) 12

ulrichi (Metopina) 14, 26,27 umbrimargo (Beckerina) 12, 19 unicalcarata (Tripbleba) 14 unispinosa (Anevrina) 12, 18, 19 unispinosa (Tripbleba) 14 urbana (Anevrina) 12, 111, 19 urbana (Triphleba) 15

vadoni (Chonocephalus) 12 velutina (Phora) 14 venusta (Dohrniphora) 13 versicolor (Diplonevra) 13 vitrea (Triphleba) 15, 34, 36 vitripennis (Gymnoptera) 13,25 vitripennis (Gymnoptera) 13 vitripennis (pseudacteon) 14 vulgaris (Spiniphora) 14

willowsi (Dohrniphora) 13 Woodiphora 15,17,37 wymani (puliciphora) 14

81

ISBN ·(1 .. 9Ii1.ls46$~i .~·~~",~~.iQ,Ul'ilfIe,YWay'ig~.S~~cl6.~