Studies of Some Fungus Gnats (Diptera: Mycetophilidae) Including Nine Additions to the British List

Systematic Entomology (1977) 2, 67-93

Studies of some fungus gnats (Diptera: Mycetophilidae) including nine additions to the British list

PETER J. CHANDLER Weston Research Laboratories, Taplow, Maidenhead, Berks.

ABSTRACT. A key is provided to the eight species of Anatella recorded in Britain, including three species newly recorded (A.dampji Landrock; A.gibba Winnertz; A.lenis Dziedzicki). A further six species are newly recorded from Britain (Exechia sororcula Lackschewitz; Exechiopsis dumitrescae (Burghele- Balacesco); Allodia (Brachycampta) angulata Lundstrom; Mycetophila autumnalis Lundstrom; M.lubomirskii Dziedzicki; M.lunata Meigen); these and several other species are redescribed. The following nomenclatural changes are proposed. Exechia bicincta (Staeger) (= spinosa Bukowski, syn.n.); Mycetophila conj'usa Dziedzicki (=affluctata Edwards, syn.n.); Mycetophila dziedzickii n0m.n. (=obscura Dziedzicki nec Walker). Female characters of seventeen of the twenty-two British species of Exechia and eight species of Mycetophila are discussed and illustrations provided.

Introduction on females but the characters employed required revision and this is attempted as far Since the revision of Edwards (1 925b) and his as practicable. Also in the Exechiini, where supplementary notes (1941), knowledge of the generic revision of Tuomikoski ( 1966) British Mycetophilidae has advanced slowly. is now generally followed, the generic position Nineteen species have, however, been added of Allodiopsis ingeniosa Kidd was found to to the British list but five have been deleted, require discussion. bringing the total to 4 16. During examination The interpretation of some species groups of recent collections of British fungus gnats, in Mycetophila is in need of clarification but several species were discovered new to the as such groups have not been clearly defined British list although known from other parts in the Palaearctic fauna, only those including of Europe; nine of them are dealt with here, species new to the British list or involving all belonging to the sub-family Myceto- nomenclatural changes are discussed here. philinae. Material of the genera concerned in Again, it has been possible to elucidate female the British Museum (Nat. Hist.) and Hope characters where these were unknown in Department of Entomology, Oxford Uni- several species. versity Museum (referred to below by the abbreviations BMNH and HD respectively) has also been studied. Anatella Winnertz A revised key to Anatella was desirable Anatella Winnertz, 1863, 854. Type-species but this can still only be applied to males as Anatella gibba Winnertz, designated by so few females have been associated. In Johannsen, 1909: 90. Exechia, more work had been done previously Correspondence: Mr P. J. Chandler, Weston Anatella species are small (1.5-3 mm in Research Laboratories, 644 Bath Road, Taplow, length), dark coloured and possess few useful Maidenhead, Berks. SL6 OPA. characters other than in the male genitalia. 68 Peter J. Chandler

Anatella is the only Palaearctic genus of suggested further specific characters, em- Exechiini to have the costa distinctly pro- ployed in the following key, which it is hoped duced beyond the tip of vein R5 and it was will be a useful aid to sorting species but considered monophyletic by Tuomikoski examination of the genitalia is still essential (1 966), who studied fifteen species. The single for confirmation. Structural characters are dark basal bristle on the hind coxa character- given precedence where possible but in some istic of all other Exechiini may be absent but instances it is necessary to place greater there are intermediate species with a weakly reliance on colour, the three types of thoracic developed bristle. The base of the cubital fork coloration being perhaps most reliable. The is usually a little beyond that of the median first flagellar segment is elongate in all species, fork (a little before in A.gibba) but its the next few being progressively shorter and position is variable within species and cannot relative measurements cited apply best to the be used for reliable identification. The fourth and immediately following segments. variation in the development of the mid tibial Although only one or two specimens have spurs mentioned by Tuomikoski and the been seen of the three additional species, they distinctive mid femoral chaetotaxy of some are easily distinguished by the highly specific spe;ies appear to be secondary sexual genital characters. These are figured in detail characters of the male. for these species but an internal view of the Eight species have been recorded from clasper complex is figured for all British Britain but three others can now be added. species except lenis. The nomenclature Only seven have been figured in British adopted has, however, required careful con- literature (setigera Edwards and unguigera sideration of the literature in the cases of Edwards in Edwards, 1921, claspers only; gibba and dan~pfi. the others in more detail in Edwards, 1925b) while A.turi Dziedzicki was added by Edwards (1941) without a figure. Eight or nine other Key to British species (males only) European species are recognized but several 1 Mid femora with posteroventral fringe well are of uncertain status. Some were figured developed, distinctly longer and stronger than by Dziedzicki (1923) and Landrock (1924, anteroventral fringe, with some hairs at least 1925) but the three described by Strobl have nearly half median femoral depth...... 2

not been figured. Strobl's rufithorax was - Mid femora with weak fringes, posteroventral founded on thoracic coloration, i.e. distinct never better developed than anteroventral, stripes on a paler ground and the type was a although some hairs of latter may be half median femoral depth ...... 4 female although he recognized the male later. 2 Mid femoral fringe strong, longest towards base I have found that this pattern may occur in where it reaches median femoral depth. Outer several species and the record by Matile (anterior) spur of mid tibia very short, not more (1967) of female rufithorax from the than one quarter length of inner spur. Antenna1 Pyrenees is therefore uncertain. flagellar segments more than twice as long as No external characters have been found in broad. No bristle at base of hind coxa. (Thorax grey pruinose; tergites 1-2 sometimes females to associate then1 with their respective slightly paler. First flagellar segment yellow males but there are probably good characters basally. Wing length 2.3-2.5 mm.) in the ovipositor. This was figured by Dzied- ciliata Winnertz zicki (1923) for A.longisetosa Dziedzicki and - Mid femoral fringe weaker, scarcely exceeding by Plassn~ann(1970b) for A.lenis Dziedzicki, half depth of femur. Outer spur of mid tibia at least half length of inner spur...... 3 of which he had reared both sexes from the 3 Flagellar segments less than twice as long as bark encrusting fungus Exidia glandulosa broad. Mid femoral fringe longer on basal half. Fries (Tremellinaceae). The rearing of A well developed bristle at base of hind coxa. A.j7avomaculata Edwards from a small (Body blackish grey. Flagellum entirely grey. lignicolous Ascomycete, Helotium aciculare Wing length 2.0-2.3 mm.) longisetosa Dziedzicki Persoon has enabled the female of this species - Ragellar segments at least twice as long as to be recognized. Nothing else is known of the broad. Mid femoral fringe longest apically but development of Anatella. not quite half femoral depth. Hind coxal bristle The present study of male Anatella has small and weak. Studies of some British Mycetophilidae 69

FIGS. 1-10. Anatella males, internal view of left claspers: (1) unguigera Edwards; (2) gibba Winnertz; (3) longisetosa Dziedzicki; (4) setigera Edwards; (5) turi Dziedzicki; (6) minuta Staeger; (7) flavomaculata Edwards; (8) dampfi Landrock; (9) simpatica Dziedzicki; (10) ciliata Winnertz.

(Body grey pruinose. First flagellar segment - Bristle at base of hind coxa usually well de- yellow basally) ...... setigera Edwards veloped, always present. Outer mid tibial spur 4 Outer spur of mid tibia absent although inner usually shorter (except in dampfi). Wing veins spur of normal strength. Flagellar segments grey, fork veins as usual paler than the radial longer than broad but not twice breadth. No sector...... 6 bristle at base of hind coxa. Mid femoral Fringes 6 Elagellar segments 3-4 about 1.5 X long as very short. broad. Outer mid tibial spur about 0.8 length of (Thorax grey pruinose; tergites 1-3 yellow inner spur. Hind coxal bristle well developed. laterally. First one or two flagellar segments Anteroventral mid femoral fringe well developed yellow. Wing length 1.9 -2.2 mm.) but less than half femoral depth. ruri Dziedzicki (Thorax grey pruinose. Base of first flagellar Outer spur of mid tibia always present; although segment pale. Wing length 3.1 mm.) almost invariably shorter than inner spur, never gibba Winnertz less than half as long ...... 5 - Flagellar segments 3-4 quadrate or a little longer .7 S Bristle at base of hind coxa absent. Outer mid than broad...... tibial spur more than five-sixths length of inner 7 Thorax uniformly sooty black on dorsum. Sides spur. Wing veins yellow. Flagellar segments from of first three tergites usually broadly yellowish the fourth onwards nearly quadrate (segment 3, but this coloration sometimes obscured or re- 1.2 X breadth). Mid femoral fringes short, duced in extent. Antennae and palpi grey, base longest apically. of first flagellar segment brownish ...... 8 (Body dark grey, thorax blackish dorsally. Basal - Thorax obscurely grey brown, often more antenna1 segments and palpi darker than in reddish brown laterally, never uniformly black A.setigera. Wing length 2.4-2 .5 mm.) but sometimes with pale ground bearing a dark ur~guigeraEdwards central stripe and shorter more or less fused 70 Peter J. Chandler

lateral stripes, leaving pale shoulders. Mid femoral anteriorly. Pleura largely pale brownish fringes short and weak, longer apically ...... 9 yellow (the thoracic coloration may display 8 Mid femora with anteroventral fringe well de- the same range of variation found in minuta veloped but short, longest apically. Outer mid tibial spur five-sixths length of inner spur. Bristle and simpatica). Halteres pale yellow. Abdo- at base of hind coxa small and weak. Wing length men grey with tergites 2-3 narrowly pale 2.3 mm...... dampfi Landrock laterally. Hypopygium brownish yellow (Figs. - Mid femora with anteroventral fringe weak and 13 and 14). not closeset but on basal half may exceed half Legs yellow. Fore metatarsus four-fifths median femoral depth. Outer mid tibial spur length of its tibia. Mid femora with fringes about three-quarters length of inner spur. Bristle at base of hind coxa well developed. Wing length very short, stronger and longer towards tip. 1.9-2.0 mm...... f[avomaculata Edwards Outer mid tibial spur two-thirds length of 9 Basal antennal segments (scape and pedicel) and inner spur. Bristle at base of hind coxa strong. palpi obscurely yellowish brown. Abdominal Wings with fork veins pale grey but tergites 1-4 with yellow patches towards their distinct. Costa extending about halfway from lower posterior corners, sometimes faint. Outer mid tibial spur of variable length, from little R5 to M1. Cubital fork beginning slightly more than half to three-quarters length of inner beyond base of median fork. Wing length spur. Wing length 1.6-2.4 mm. 2.4 mm. simpatica Dziedzicki Material examined. HUNTS: Monk's Wood, - Basal antennal segments and palpi yellow but 15 .ix. 1972, 1 6 ex suction trap (Cole). antennal flagellum usually entirely grey. Outer mid tibial spur about two-thirds as long as inner. 10 Anatella gibba W innertz 10 Flagellar segments slightly longer than broad. Fork veins pale grey but more distinct than in Anatella gibba Winnertz, 1863, 855; Dzied- minura. Thorax dull orange brown with three darker stripes. Wing length 2.4 mm. zicki, 1923: 3, Figs. 11 and 12 (based on lenis Dziedzicki Winnertz' type). - Flagellar segments quadrate or slightly broader Landrock's (1927) character of the base of than long. Fork veins very pale. Coloration variable, thorax often all dark grey brown but the cubital fork being a little before that of usually paler laterally and may have colour the median applies to the British specimen described for lenis. Sides of tergites 2-3 some- and may be constant. 1 did not use it in the times pale. Wing length 1 .8 -2 0 mm. key because I found that some other species mirzuta Staeger may have it either immediately below or distinctly beyond the base of the median fork. Landrock did not reproduce a figure of Anatella lenis Dziedzicki the genitalia because he was then unaware of the work of Dziedzicki (1923) (published Anatella lenis Dziedzicki, 1923; 5, Figs. 23- posthumously, the author having died in 24. 1921), who figured dorsal and ventral views. The British example agrees well with Dzied- The British example agrees with these figures zicki's (1923) figures and there is little doubt in the slender lower claspers, hairy only at the that it is correctly identified. A.lenis was tip and the nearly equally bilobed broad recorded by Lackschewitz (1937) from the posterior margin of the upper claspers. It Baltic States and Plassmann (1970a, b, 1971, apparently differs in the relative dimensions 1973), who reared it in Germany, described of the median and lateral sterna1 processes the larva and figured the ovipositor but in a but this may be due to the angle at which contracted position. they were drawn (in Dziedzicki's figure the Male. Head grey. Palpi and basal antennal lateral processes are much shorter). segments yellow; flagellum dark grey. Male. Head dark grey. Antennae with basal Flagellar segments slightly longer than broad. segments dark yellowish brown, flagellum Thorax dull orange brown with three darker grey with base of first segment paler. Palpi stripes, the median broadened anteriorly to grey brown. Flagellar segments 3-4 1.5 the fore margin, the laterals ending where the times as long as broad. Thoracic dorsun? grey median begins to broaden and rounded pruinose, slightly paler laterally. Pleura grey Studies of some British Mycetophilidae 7 1

FIGS. 11 12.Anatella gibba Winnertz, rnale genitalia: (l l) ventral view; (12) dorsaI view. but propleura brownish yellow. Halteres flanges articulating with the genital capsule brownish with paler stalk. Abdomen grey. but not very exactly in their shape. His Hypopygium brownish yellow (Figs. l 1 and lateral view of the claspers, however, is con- 12; claspers, Fig. 2). sidered confirmatory as their shape is nearly Legs yellow. Fore metatarsus equal to tibia identical; the thin menibranous internal in length. Mid feinoral fringes short, anterior process is omitted but its shape and position fringe increasing in length apically where it are indicated in his dorsal view. approaches half median femoral width. Mid Landrock (1937) suggested that A.novata tibia with outer spur three-quarters length of Dziedzicki (1923) might be the same as inner. A welldeveloped bristle at base of hind darnpfi but noilata is not used here because coxa. in his figures of novata the upper claspers re- Wing veins yellowish grey, fork veins semble the lower claspers of danzpfi' from the paler. Costa extending half distance from K5 views figured and I think this synonymy un- to M1. Cubital fork begins slightly before likely. According to Landrock's (1927) key, l base of median fork. Wing length 3.1 mm A.darnpfi (like A.unguigera Edwards) has the (the largest specimen of Anatella examined outer mid tibial spur nearly as long as the but it may be abnormally large for the inner as in the British specimen. The dark species). thoracic coloration is also found in unguigera, Material examined. INVERNESS: Kinrara, fZavomaculata and longisetosa but the 23.vi. 1967, l d (D. M. Ackiand, BMNH). genitalic structure is nearer to sirnpatica. Male. Head grey dusted. Antennae and palpi grey, but basal antenna1 segments and first flagellar segment brownish. Flagcllar Anatella darnpfi Landrock segments nearly quadrate, slightly longer than broad. Thoracic dorsum slightly shining Anatella darnpfi Landrock, 1924: 80, Figs. sooty black; pleura and metanoturn brownish 8-10. grey. Halteres yellow. Abdomen dark grey Landrock (19 24) described A.dampfi from with tergites 2-4 yellowish laterally. Hypo- Estonia and figured the genitalia. If his dorsal pygiurn small, brownish yellow (Figs. l S and and ventral views represent the species des- 16; claspers, Fig. 8). cribed here they must be inaccurate, agreeing Legs yellow. Fore metatarsus equal to in the lateral sterna1 processes being separate tibia. Mid femoral fringes short and weak. 72 Peter J. Chandler

Mid tibia with outer spur five-sixths length of Claspers, Fig. 1; 128 seen from Britain inner spur. Hind tibia (as usual) with outer (Sussex to Sutherland) and Ireldnd (Down). spur three-quarters length of inner. A small weak bristle near base of hind coxa. Anatella longisetosa Dziedzicki Wings with grey veins, fork veins paler. Costa extending two-fifths of distance from Anatella longisetosa Dziedzicki, 1923: 6, R5 to M1. Base of cubital fork distinctly Figs. 27-31. beyond base of median fork. Wing Iength Anatella piligera Edwards, 1925b: 590, Figs. 2- 3 mm. 48-50 (examined). Material examined. MERIONETH: Cwm Claspers, Fig. 3; 16 d seen from Britain Bychan, 12.x. 1975, l d in hillside oakwood (Bucks. to Inverness). (Ismay); SELKIRK: Happertutie Burn, iv. 1976, l d (B. Milligan). Anatella setigera Edwards

Anatella setigera Edwards, 192 1 : 122, Fig. 1. Anatella unguigera Edwards Claspers, Fig. 4; 14 d seen from Britain Anatella unguigera Edwards, 1921 : 122, (Devon and Hunts. to Argyllshire) and Ireland Fig. 2 (examined). (Sligo).

FIGS. 13-16. Anatella male genitalia: (1 3-14) lenis Dziedzicki, dorsal and ventral views; (15-1 6) dampfi Landrock, dorsal and ventral views. Studies of some British Mycetophilidae 73

Anatella turi Dziedzicki Exechia Winnertz Anatella turi Dziedzicki, 1923: 4, Figs. 17 Exechia Winnertz, 1863: 879. Type-species and 18. Mycetaphila fungorum Degeer, designated by Johannsen, 1909: 106 (rnisident.)= Claspers, Fig. 5; 10 d seen of this previously fusca (Meigen). little known but evidently widespread species. New records are: HUNTS: Monk's Wood, This name is restricted, following Tuomi- 28.vi.1971, 4d, 25.viii.1972, l d (Cole); koski (1966) to Group I of Edwards'(1925b) CAMBS: Chippenham Fen, 24 .ix. 194 1, d key, with twenty-one British species. Many (Collin, HD); CAERNARVON: Glan-y-Gors, are only reliably determined by reference to 16 .vii. 1976, d; ROSS: Kinrive pinewood, the male genitalia and females have been 15 .vi. 1976, 2 d (Chandler). inadequately studied. The discovery of both sexes of an Exechia new to Britain led to a re-assessment of the characters used to Anatella minuta (Staeger) distinguish species of the bicincta Staeger- lundstroemi Landrock Group, to which it Mycetophila minuta Staeger, 1840: 253 belongs. Females of other species were studied (examined by Edwards, 1925a). and good characters in the ovipositor were Anatella minuta (Staeger): Edwards, 1925a, found although differences are less pro- 167. nounced between related species than in the Claspers, Fig. 6; l l d seen, mostly from male genitalia. southern England and Wales but one from Many published figures of Mycetophilid Scotland (INVERNESS: Inverdruie). ovipositors show the contracted position, often concealing important features, while inaccuracy and misidentification are some- Anatella flavomaculata Edwards times involved. In Exechia, the shape of the pregenital tergite (VII), sometimes omitted Anatella flavomaculata Edwards, 1925b: 590, from figures, is highly specific and often Figs. 5 1- 53 (examined). permits identification of dried material. As Claspers, Fig. 7; ovipositor, Figs. 17 and 18; few ovipositors have been figured in sufficient 248 and 19 seen from many parts of Britain. detail for accurate recognition, those which Two males and one female were reared by have been identified in the material examined R. E. Evans from Helotiwn aciculare collected are discussed and figured. at Oversley Wood, WARWICKS. in ix. 1972.

Exechia sororcula Lackschewitz Anatella simpatica Dziedzicki Exechia sororcula Lackschcwitz, 1937: 22, Anatella simpatica Dziedzicki, 1923: 6, Figs. 6a-g. Figs. 25 and 26. This little known species was frequent in the Anatella incisurata Edwards, 1925b: 589, Dolgellau area of north Wales in October Figs. 43-45 (examined). 1975. Apart from Lackschewitz' unique Claspers, Fig. 9; 24 d seen from Britain Estonian type, no records have appeared (Sussex to Sutherland). although it was included in the keys of Landrock (1940) and Ostroverkhova & Stackelberg (1969). It runs in the key of Anatella ciliata W innertz Edwards (1925b) to E.lundstroemi Landrock, of which only one British male is certainly Anatella ciliata Winnertz, 1863: 856; Dzied- known (Edwards, op. cit.) although it is said zicki, 1923: Figs. 13 and 14 (based on to be not uncommon in some parts of Europe. Winnertz' type). The locality for lundstroemi was not re- Claspers, Fig. 10; 3 1 d from all parts of the corded, only that it was in Mr Collin's collec- British Isles. tion; I have not, however, located the speci- 74 Peter J. Chandler

t'IGS. 17-20. Female genitalia of Anatella and Exechia: Anatella f7avomaculata Edwards, (1 7) lateral view; (18) dorsal view; Exectlia sp. near dizona Edwards (?cincta Winnertz), (19) lateral view; (20) abdominal pattern.

men. Comparison of sororcula with bicinctu patches and yellowish propleura. Two pro- Staeger is also necessary because this often pleural bristles. Abdomen with tergites 1 runs to lundstroemi in keys. and 6 entirely dark, 2-5 with yellow lateral The hypopygium of sororcula is distinctive; spots in contact with fore margins, that on the external claspers are rounded and bear 2 elongate, those on 3-5 more rounded, all long dense apical bristling. There are yellow occupying at least half tergal length, narrowly markings on the bases of the male tergytes but distinctly separated dorsally. Hypopygium 2-5 compared with 2-4 in bicincta and brownish yellow (Figs. 2 1-23). Legs yellow only 3-4 in lundstroenli. The female soror- with distinct dark patches beneath bases of cula, not previously recognized, has yellow mid and hind femora. Hind tibia with 3-5 d, markings on tergites 3-6 while bicirlcta has 6-10 a, 4-6 short p near tip. them on 2-5; the female characters of lund- Female. Very similar to male except in stroerni require clarification. The ovipositor abdominal pattern. Tergites 1-2 entirely is obviously different from' bicincta as in- dark; 3-6 have large yellow lateral spots at terpreted below. That of lundstroerni has bases, those on 3-4 not in contact with fore only been figured by Ostroverkhova & margin on upper half, 5-6 only touching fore Stackelberg (op. cit.) but the figure may margins at lower corners, that on 6 distinctly represent bicincta which it resembles. smaller than those on 3-5. Tergite 7 and Male. Head dark, with yellow palpi and ovipositor (Fig. 24) brownish yellow. basal antenna1 segments; flagellum dark Wing length 3.25 mm. except basal half of first segment. Thorax Material examined. MERIONETH: largely dark with small yellow humeral Brithdir-Isaf, 25 .ix.197 1, d, 9 (Hutson, 76 Peter J. Chandler

Even where in the male there appear to be basal bands on tergites 2-4 of the male bands in lateral view, they are usually occupying a little more than half the tergal narrowly divided dorsally. There is also length dorsally and extended to the apical variation in the number of anterior (external) margin laterally; there may be a constant bristles on the hind tibia (normally 12-14 difference in this pattern. Only two males of in a close set row, but ranging from 9 to 15). dizona have been recorded from Britain, one The ovipositor of E.bicincta (Fig. 25) is from the Clifton collection, locality un- easily recognized; the figures of Lundstrom known, in BMNH, the other (DORSET: (1 9 11) and Dziedzicki (1 91 5) fit the British Studland, 5 .X.1910, J. W. Yerbury) I found specimens and the figure of spinosa Bukowski unnamed in the Verrall-Collin collection, by Matile (1963) also agrees quite well. In- HD. Edwards (1925b) mentioned a female deed, the figures of male genitalia given by from Monk's Wood, HUNTS., but I found Bukowski (1949) and Matile (op. cit.) for that it is one of the possible repanda spinosa also agree with bicincta. I believe that discussed below. The true female of E.dizona spinosa was based on specimens of bicincta has not yet been recognized but a French with incomplete abdominal bands, leading it female, briefly described below, might be to be compared only with lundstroerni; dizona or more likely belongs to the non- Dr L. Matile tells me that he is now also of British cincta Winnertz of which the female this opinion. has not been described. It runs to the The figure of the ovipositor of lundstroemi bicinctaldizona couplet in Edwards' key and (Ostroverkhova & Stackelberg, 1969) agrees is distinct from bicincta in both abdominal with bicincta in the lack of a clearly pattern and ovipositor; it has a very character- differentiated apical cercal segment, present istic pregenital tergite. in all other Exechia 1 have examined. Al- Female. Head grey dusted. Antennae with though there are some small differences, it is basal segments and base of first flagellar possible that this too is based on bicincta. segment yellow, rest grey. Palpi yellow. The only published British records of Thorax mainly dusted dark grey, yellow on bicincta 1 have found are by Edwards (1925b, shoulders and narrowly on fore and lateral 1933) and Kidd & Ackland (1970). Only margins of mesoscutum. Abdomen with males were present in the BMNH collection complete yellow bands basally on tergites and a female labelled ?bicincta teste Edwards 3-6; abdominal pattern and ovipositor, in HD was B.dorsalis (Staeger). My identifica- Figs. 19 and 20. Legs yellow with dark tion of the female is based on a series (3 6, patches beneath bases of mid and hind 3 Q) I reared from an unidentified small pale femora. Two strong propleurals, with a short brown gill fungus (collected in a copse at weak one behind. Hind tibia with 7 8 d, Bromley, KENT, 19.v. 1965, pupated 22.v., 8-10 a, 4-6 short p near tip. emerged 29 .v.). I have in all examined 16 6 Material examined. FRANCE (Charentes- and 69, from southern England north to Maritimes): Saintes, 23. v. 197 1, Q swept on a Suffolk and Hereford except for two females wooded roadside verge (Chandler). from north Wales which have the thorax mainly dark greyish instead of the usual coloration where it is broadly yellow laterally The females of other Exechia species and brownish dorsally. Females have been associated as far as possible with their males in the remaining British Execlzia dizona Edwards species of the genus. Some conclusions are tentative but it is hoped that this preliminary [Exechia bicincta (Staeger) Lundstrom, 1909: work will be a stimulus to further study based 46, Figs. 94 and 95. Misident.] on reared series and pairs taken in copula. Exechia dizona Edwards, 1925a: 166 (n0m.n. for bicincta Lundstrom nec Staeger). Execlzia seriata (Meigen ) This species, coupled with bicincta in Edwards' ( 1925b) key, has complete yellow Exechia seriata (Meigen); Edwards, 1924: 16. Studies ofsorne British Mycetophilidae 75

FIGS. 2125. Genitalia of Exechia: sororcula Lackschewitz, (21) male hypopygium, ventral view; (22) dorsal view; (23) eighth sternite; (24) ovipositor, lateral view; bicincta Staeger, (25) ovipositor, lateral view.

BMNH); Benglog, 9.x. 1975, 0; near 1863: 895; Dziedzicki, 1915: Tablc XV, Mallwyd, 12.x. 1975, 3 d; Cwm Llinau, Figs. 236 and 237 d, Table XVI, Figs. 238 12.x.1975, 3 d (Stubbs); Coed Ganllwyd, and 239 ? (based on Winnertz' material); 10.x. 1975, 9 (Stubbs), d (Hutson, BMNH); Edwards, 1913: 371. Ceunant Llannyrch, 11 .X.1975, d (Stubbs), Execliia serpentino Lundstr6m, 191 1: 407, 128 (Hutson, BMNH); Coed Crafnant, Taf. XIII, Figs. 9-12. dQ. 12.x.1975, d (Hutson, BMNH); Cocd y Exechio bicincto (Staeger); Edwards, 1925a: Brenin, 25 .X.1975, 3 d (Chandler); CAER- 168; Edwards, 1925b: 594. NARVON: Coed Trcmadog, 11 .X.1975, d Exechia spinosa Bukowski, 1949: 407, Fig. 5. (Chandler) (all localities were in more or less d. Syn.nov. mixed deciduous woodland). Exechia spinosa Bukowski; Matile, 1963: 141, Figs. 1 and 2.89. The record of E.lundstroerni which 1 Exechia bicincta (Staeger) published (1966) was based on a female of hfycetophilo bicincta Staeger, 1840: 263; bicincta. Because of itsinconlpleteabdominal (examined) Edwards, 1925a. bands, it ran to lundstroerni in Edwards' Mvcetophila interrupt4 Zetterstedt, 185 2: key but examination of other material of 4240; (examined) Edwards, 1925a. bicincta has shown that the female has Execlzio interrupts (Zetterstedt); Winnertz, narrowly separated lunules rather than bands. 76 Peter J. Chandler

Even where in the male there appear to be basal bands on tergites 2-4 of the male bands in lateral view, they are usually occupying a little inore than half the tergal narrowly divided dorsally. There is also length dorsally and extended to the apical variation in the number of anterior (external) margin laterally; there may be a constant bristles on the hind tibia (normally 12-14 difference in this pattern. Only two males of in a close set row, but ranging from 9 to 15). dizona have been recorded from Britain, one The ovipositor of B.bicincta (Fig. 25) is from the Clifton collection, locality un- easily recognized ; the figures of Lundstrom known, in BMNH, the other (DORSET: (191 1) and Dziedzicki (1915) fit the British Studland, 5 .X.19 10, J. W. Yerbury) I found specimens and the figure of spinosa Bukowski unnamed in the Verrall-Collin collection, by Matile (1963) also agrees quite well. In- HD. Edwards (1925b) mentioned a female deed, the figures of male genitalia given by from Monk's Wood, HUNTS., but I found Bukowski (1949) and Matile (op. cif.) for that it is one of the possible repanda spinosa also agree with bicincta. I believe that discussed below. The true female of E.dizona spinosa was based on specimens of biclncta has not yet been recognized but a French with incomplete abdominal bands, leading it female, briefly described below, might be to be compared only with lundstroerni; dizona or more likely belongs to the non- Dr L. Matile tells me that he is now also of British cincta Winnertz of which the female this opinion. has not been described. It runs to the The figure of the ovipositor of lundstroerni bicinctaldizona couplet in Edwards' key and (Ostroverkhova & Stackelberg, 1969) agrees is distinct from bicincta in both abdominal with bicincta in the lack of a clearly pattern and ovipositor; it has a very character- differentiated apical cercal segment, present istic pregenital tergite. in all other Exechia 1 have examined. Al- Female. Head grey dusted. Antennae with though there are some small differences, it is basal segments and base of first flagellar possible that this too is based on bicincta. segment yellow, rest grey. Palpi yellow. The only published British records of Thorax mainly dusted dark grey, yellow on bicincta I have found are by Edwards (1 925b, shoulders and narrowly on fore and lateral 1933) and Kidd & Ackland (1970). Only margins of mesoscutum. Abdomen with males were present in the BMNH collection complete yzllow bands basally on tergites and a female labelled ?bicincta teste Edwards 3-6 ; abdominal pattern and ovipositor, in HD was Edorsalis (Staeger). My identifica- Figs. 19 and 20. Legs yellow with dark tion of the female is based on a series (3 d, patches beneath bases of mid and hind 3 9) I reared from an unidentified small pale femora. Two strong propleurals, with a short brown gill fungus (collected in a copse at weak one behind. Hind tibia with 7 8 d, Bromley, KENT, 19 .v. 1965, pupated 22.v., 8-10 a, 4-6 short p near tip. emerged 29.v.). I have in all examined 166 Material examined. FRANCE (Charentes- and 69, from southern England north to Maritirnes): Saintes, 23 .v. 197 1, 9 swept on a Suffolk and Hereford except for two females wooded roadside verge (Chandler). from north Wales which have the thorax mainly dark greyish instead of the usual coloration where it is broadly yellow laterally The females of other Exechia species and brownish dorsally. Females have been associated as far as possible with their males in the remaining British Exechia dizona Edwards species of the genus. Some conclusions are tentative but it is hoped that this preliminary [Exechia bicincta (Staeger) Lundstrom, 1909: work will be a stimulus to further study based 46, Figs. 94 and 95. Misident.] on reared series and pairs taken in copula. Exechia dizona Edwards, 1925a: 166 (n0m.n. for bicincta Lundstrom nec Staeger). Exechia seriata (Meigen) This species, coupled with bicincta in Edwards' ( 1925b) key, has complete yellow Exechia seriata (Meigen); Edwards, 1924: 16. Studies of some British Mycetophilidae 77

Exechia pallida (Stannius); Lundstrom, 1909: Exechia lucidula (Zetterstedt) Figs. 126 and 127; Edwards, 1925b: 593 (apparently a lapsus). This is the only British Exechia with a strongly shining black thorax, also found in The ovipositor, figured by Lundstrom (1909), two other Palaearctic species, nitidicollis is dorsoventrally flattened unlike other Lundstrom and micans Lagtovka & Matile. Exechia I have seen; combined with the other The ovipositors of these last two were figured specific characters it is easily recognized. by Lagtovka & Matile (1 974); E.lucidula,(Fig. 27) resembles them but is distinct, being closest to micans in the small apical cercal Exechia dorsalis (Staeger) segment. E.1ucidula has not apparently been col- Exechia dorsalis (Staeger); Dziedzicki, 1915: lected in Britain since 194 1 ; it is known from Figs. 25 1 and 252. HERTS., SUFFOLK (Edwards, 1913, 1925b), Ovipositor, Fig. 26. The scalloped margin of CHESHIRE (Kidd, 1959) and CAMBS. (Chip- the pregenital tergite is distinctive. penham Fen, 1 d, 20 .vii.1941, Collin, HD).

FIGS. 26-29. Ovipositors of Exechia species, in IateraI view: (26) dorsalis Staeger; (27) lucidula Zetterstedt; (28) fusca Meigen; (29) confinis Winnertz. 78 Peter J. Chandler

Exechia spirluligera Lundstrom Exechia festiva Winnertz Exechia spinuligera Lundstrom, 19 12: 33, E.~echia festiva Winnertz; Lundstrom, 1909: n0m.n. for spinigera (Winnertz) Lund- 43, Figs. 121 and 122. strotn, 1909: 45, Figs. l l l and 112. Ovipositor, Fig. 32. Lundstrom's (1909) [Exechia spinigera (Winnertz); Edwards, figure is probably correct but shows the 1925 ; 594. Misident.] contracted condition and omits the slight Ovipositor, Fig. 30. Lundstrom's (1 909) median apical projection of the pregenital tergite. figures agree but related species Cfvigida Boheman in Britain and three other Palae- arctic species) may be very similar. The dark Exechia pseudofestiva Lackschewitz mark beneath the base of the hind femora may be faint or absent in spinuligera (and The ovipositor figure (Fig. 33) is based on the occasionally fusca Meken), which then run female of the pair recorded by Edwards to the parva Group in Edwards' key but are (1941); there is no indication that they robust apart from the were taken in copula but the structure is differences. distinct from any other seen. Apart from the above and the recent Irish record (Chandler, 1976), I have seen 26: MERIONETH: Coed The Exechia fuscu (Meigen) Group Crafnant, 12.X. 1975, (Chandler); INVEK- NESS: Ben Macdhui, 15 .viii. 1968. pit-fall The ovipositors of the two British species, trap at 3870 ft (R, C, Welch). fusca (Meigen) (Fig. 28) and confinis Win- nertz (Fig. 79) have sorlle small differences basally which appear to be constant. Previous Exechia separata LundstrGm figures of fusca (Lundstrom, 1909; Dzied- zicki, 1915 ; both as jungorunl D~~~~~),E.yechia separata Lundstrom, 19 12: 34, conji,zis (Plassmann, 1970b) and peyerim- n0m.n. for cotzcinna (Winnertz) Lurid- hoffi Burghele-Balacesco ( 1966) are in- strom, 1909, 46, Figs. 132 and 133 (?), separable. 87-89 (6). [Exechia lateralis (Meigen); Dziedzicki, 19 15, Figs. 255 and 256, 0. Misident.] Exechia nigra Edwards Dziedzicki (19 15) figured dorsal and ventral views, showing a broad emargination of the Exechia tzigra Edwards, 1925b: 595, Fig. 63a. tergite but omitting the long fine The shape of the pregenital tergite is dis- marginal hairs. Lundstrom's (1909) figure tinctive but Plassrnann (1970) figured the agrees in the short cerci but shows only a non-British nigrofusca Lundstrom, showing no shallow emargination. The female figured here obvious difference frorn nigra although they (Figs. 34 and 35) has been reared from are not closely allied. Russula species with males of E.separata.

The Exechia contarnirlata Winnertz Group Exechia parva Lundstrom

E. con taminata (Fig. 3 1 ) was correctly British specimens agreeing with Plassmann's figured by Lundstrom (1909, as dorsalis) (1970) figure of parva have two types of and Dziedzicki (1 9 15). Some examples which abdominal marking. The first group, con- agree, however, run to nigroscutellata Land- sidered to be typical parva (Fig. 36), have rock in keys because they lack a dark mark small yellow patches on tergites 3-6 not beneath the hind femora. The ovipositor of more than half tergal height in lateral view. nigroscutellata (Ostroverkhova & Stackelberg, The second group have a yellow band on the 1969) is apparently distinct but the female of side margin of tergite 2, yellow patches pseudocincta Strobl has not been recognized. extended dorsally as complete or narrowly Studies of some British Mycetophilidae 79

FIGS. 30-33. Ovipositors of Exechia species, in lateral view: (30) spinuligera Lundstrom; (31) contamirrata Winnertz; (32) frstiva Winnertz; (33) pseudofestiva Lackschewitz.

divided narrow basal bands on 3-4 (longest 42, Figs. 117 and 118, form A; Fig. 119, on 4, two-fifths tergal length), triangular form B. Misident.] patches on 5-6 and 7 yellow but vaguely Exechia nana (Staeger); Edwards, 1925a: darkened dorsally. These run in Edwards' 167. key to esigua Lundstrom but Ostroverkhova Lundstrom figured two forms of the ovi- & Stackelberg (1969) figured exigua with positor, both with elongate cerci, one with a longer cerci and resembling the probable produced apical margin to the pregenital nana discussed below. Also some were found tergite, the other with a central emargination. hibernating in unibelliferous stems at Clumber The first (Fig. 37) was identified as nana by Park, NOTTS. with males of repanda Edwards (4P in BMNH: Baldock, HERTS.; Johannsen and they may belong to this Shefford, BEDS.) but only one other has species which has Inale genitalia close to been seen (SURREY: Thursley Common, parva. 16.iv. 1967, A. E. Stubbs). They have large yellow lateral patches on tergtes 3-4 (two- thirds tergal height) and smaller patches on Execlria nana (Staeger) 2, 5 and 6. A commoner British form (8 0; [Esechia lateralis (Meigen); Lundstrom, 1909: KENT, SURREY, MERIONETH and 80 Peter J. Chandler

FIGS. 34-38. Ovipositors of Exechia species: (34) separata Lundstrom, lateral view, (35) dorsal view of pregenital tergite; (36) parva Lundstrom; (37) ? exigua Lundstrom; (38) ? nanu Staeger.

ARGYLLSHIRE) (Fig. 38) with only small separable on genital characters from intersecta yellow lateral spots on the tergites, could be Meigen and magnicauda Lundstrom, may Lundstrom's second form, which he suggested be added. My record (1966) of magnicauda might be parva. Ostroverkhova & Stackel- from Kent was incorrect, being based on berg's (1969) figure of exigua was similar intersecta. but 1 consider it more probable that it is the true nana, widespread in Britain, while the first form could feasibly be exigua, a local Exechiopsis (E.) dumitrescae (Burghele- southern species recorded from Baldock and Balacesco) Shefford by Edwards (1 925 b). C'xechia dumitrescui Burghele-Balacesco, 1972: 390, Plate 124: 3. Exechiopsis (E.) dumitrescae (Burghele- Exechiopsis Tuomi koski Balacesco): LaStovka & Matile, 1974, 1 13, Exechiopsis Tuomikoski, 1966: 177. Type- Fig. 38. species Exechia subulata Winnertz, by Desc~iked from three males collected in a original designation. Rumanian cave, one male has since been Most of Edwards' (1925b) Group I1 of found in Mongolia (LaStovka & Matile, 1974). Exechia are now placed here, comprising I have examined one male which agrees with fifteen British species. A further species, only the published figures and its collector tells Studies of some British Mycetophilidae 8 1

FIGS. 39-40. Exechiopsis dumitrescae Burghele-Balacesco, male hypopygium: (39) dorsal view; (40) ventral view. me that he has four further males from the laterally to bases of tergites 1-3, more same locality (28 .vi. 1971, 18 .vii. 1971, restricted on 4-5; 6 only narrowly yellow on 26.ix. 1972, 29.ix. 1972). hind margin. Hypopygium brownish yellow Male. Head brownish. Antennae with basal (Figs. 39 and 40). Wings clear. Sc ends in R. joints and basal half of first flagellar segment R5 strongly downcurved near tip. Petiole of yellow, the rest darkened. Thorax mainly dull m-fork about half length of r-m. Legs yellow; yellow with three discrete light brown stripes fore metatarsus about one sixth longer than dorsally, a long median reaching the fore its tibia. Hind tibia has 11 -1 2 a, 9 d, 10-1 1 margin but tapered posteriorly to only faintly short p near tip. Wing length 4 mm. reach the scutellum and short laterals leaving Material examined. HUNTS: Monk's Wood, broadly yellow humeral areas and sides; dark 10 .vii. 1972, d ex suction trap (Cole). median stripe on scutellum and metanotum; some darker shades on lower parts of pleura. Allodia Winnertz Discal bristles well developed on '.Jm soscutuin; one long propleural -(t- a second sub-genus Brachycampta Winnertz short hair-like propleural~ Tergites largely Brachycampta Winnertz, 1863: 833 ; Tuomi- yellow posteriorly, the yellow extending koski, 1966: 184. Type-species hlyceto- 82 Peter J. Chandler

phila alternans (Zetterstedt) Winnertz Allodia (Brachycampta) angulata (Lundstrom) (= Allodia grata (Meigen) Edwards, 1925b: Brachycampta angulata Lundstrom, 1913: 607), designated by Coquillett, 1910. 308, Taf. XV, Figs. 5 and 6. Tuomikoski's (1966) concept of this sub- Allodia angulata (Lundstrom); Landrock, genus includes the grata (Meigen) and sil~~atica 1927: 119, Taf. lX, Fig. 21. (Landrock) Groups of Edwards' (1925 b) key. One British male accords with Lundstrom's It is characterized by well-developed discal figures of angulata, described from one male bristles on the mesoscutum, yellow abdominal collected in Swedish Lapland, 8.vii .l91 1 and markings more extended towards bases of not recorded since. Lundstrom did not tergites and a longer cubital fork, the base of mention discal bristles and Landrock (1927) which is level with the base of r-m in czernyi concluded that it lacked them. They are, (Landrock) and alternans (Zetterstedt). The however, present in the British specimen and fork is longer and distinctly before r-m in it is evident that angulata should be placed in the other species. Reliable separation of Brachycampta. species cannot be achieved from external Male. Head dark brown; palpi and base characters but the male genitalia are highly of antennae, including basal half of first 5 specific in contra#t to the close similarity flagellar segment, yellow; rest of flagellum between species of'~1lodiasensu stricto. brownish. Mesoscutum mainly brown but Recent material has been seen of the eight humeral area and continuous side margins British species and a ninth can now be added. yellow (only a small humeral spot in Type). The only common species is grata (Meigen), Scutellum brown, metanotum yellowish; which has been reared from various large or pleura brownish yellow, propleura paler medium sized terrestrial Agarics and Boleti yellow. Well developed black discal bristles on (summarized by Plassmann, 1971). It is mesoscutum. Two strong propleural (pro- closest to alternans which probably has episternal) bristles, four strong bristles on side similar habits as Kidd (1964) reared it from a of pronotum. One pair of strong scuteuars. Boletus species. Buxton (1960) recorded Halteres yellow. Abdomen dark brown with rearing silvatica and triangularis (Strobl) sides of tergites 14broadly yellow, on 3--4 from the cup fungus Peziza repanda Persoon; extended dorsally on fore margins; 5 with examination of Buxton's material (in BMNH) triangular yellow markings on fore margin has shown that most 'silvatica' were barbata nearly reaching hind margin; 6 entirely black (Lundstrom) (l d silvatica, l l d barbata, (the Type only had triangular spots on 2-4 4 d triangularis, 16 9 or lacking abdomen). but colour variation according to latitude No other Mycetophilidae have been reared and climate is common in the Exechiini). from Pezizales and this may suggest the usual Hypopygium large, yellow (Figs. 4 1-43). food-plants of the silvatica Group. Legs yellow; hind tibia with 11 a, 5 d, no p A. (B.) alternans is less common than bristles. grata, which is the only species known from Wings clear. Vein Sc short, ending in R; Ireland, but occurs throughout Britain. Of the R5 slightly downcurved apically. Petiole of other members, silvatica and pistillata (Lund- m-fork nearly equal to r-m. Base of cu-fork strom) are local in southern England and twice length of M1 +2 before base of m-fork. Wales; barbata and neglecta (Edwards) are Vein An short and weak but nearly reaching scarce but found in the south and in northern base of cu-fork. Wing length 2.5 mm. Scotland. A. (B.) czernyi was known from Material examined. SURREY: Chobham only one Scottish male but I obtained l d Common, sallow carr in a valley on heath- at Tummel Forest, PERTHSHIRE, 5.vii. 1975; land, 5 .viii. 1967, d (Chandler). this example has the abdomen darker than usual with the yellow markings reduced to small basal spots on tergites 2--4, occupying Allodiopsis (Gymnogonia) ingeniosa Kidd less than half the tergal length. A. (B.) triangularis too was known only from northern Allodiopsis (sub-genus Gymnogonia) ingeniosa Scotland but I can record l d from Windsor Kidd, 1969: 201, Figs. 1-3. Forest, BERKS., 22.vi.1971 and l d from This was described from a single male Glen Nant, ARGYLLSHIRE, 9 .vii. 1974. collected at Wytham Wood, Berks: two Studies of some British Mycetophilidae 83

FIGS. 41 -43. Allodia angulata Lundstrom, male hypopygium: (41) ventral view; (42) internal view of right clasper complex; (43) dorsal view. further examples have been seen. Kidd did not generic position of ingeniosa appears in- mention that the anal vein (An) is shorter conclusive and it may be regarded as a link and weaker than is normal in 'Rymosia' sensu between Allodiopsis and Allodia. lato of which Allodiopsis is a segregate pro- Some infraspecific variation should be posed by Tuomikoski (1966), who divided it noted. The Monk's Wood male has vein M1 into four possibly paraphyletic sub-genera. abbreviated on both wings, the apical quarter The sub-genus Gymnogonia was considered being absent as are the setulae of this vein close to Allodia but nearest to Allodia sensu although present on the complete M2. The strict0 with which it agreed in the lack of Alyth specimen has normal venation but its discal bristles. Kidd noted that A.ingeniosa abdominal coloration is darker, the pale did not entirely conform' to Gymnogon.ia markings being reduced to apical patches on because it possesses discal bristles on the tergites 2-4. posterior part of the mesoscutum. Material examined. HUNTS: Monk's Wood, In ingeniosa the anal vein only extends 29 .ix. 1972, d ex suction trap (Cole); PERTH- to two-thirds of the length of the stem of the SHIRE: Den of Alyth, 31 .vii. 1975, d swept cubital fork whereas in typical Allodiopsis it in mixed woods (Chandler). is prolonged past the base of that fork. It therefore runs in Edwards' (1925b) key to Allodia; A. (B.) czernyi has some similarities Mycetophila Meigen in the genitalia but has An very short, ending at one third of the Cu stem. The silvatica Mycetophila Meigen, 1803: 263. Type-species Group, however, have An reaching the base of Tipula agarici Villers, designated by the cubital fork because this is nearer the Johannsen, 1909. base of the wing. A.ingeniosa also agrees with This is the largest genus of Mycetophilidae Allodia in the small body size compared with with many species groups meriting closer typical robust Allodiopsis. It does, however, study. Edwards (1925b, 1941) recognized possess the posterior hind tibial bristles and 49 British species. Kidd & Ackland (1970) the setulae on the tips of the fork veins added bohemica (LaStovka), one of the characteristic of Allodiopsis and not present vittipes Zetterstedt Group of which LaStovka in Allodia. Although it agrees with Brachy- (1963) revised the European species. campta in the thoracic bristling it differs in LaStovka & Kidd (1975) recognized four the posteriorly set abdominal markings. The British members of the ruficollis Meigen 6 84 Peter J. Chandler

Group (=lineola Meigen of Edwards). Here bimaculata Fabricius) and forcipata Lund- three further Mycetophila species are re- strom, later (1926) adding tridentata Lund- corded, bringing the British list to 56. Two strom. Two further species which run to nomenclatural changes have also been found M.bimaculata in Edwards' key are here added, necessary. both on males only, their females being un- Laffoon (1 957) revised the Nearctic known. M.autumnalis Lundstrom appears to species, drawing attention to several external belong to the pictula Group and probably has characters which help to associate the sexes the usual female characters but lubomirskii where reliance had been placed almost Dziedzicki has rather different male genitalia entirely on the male genitalia. Twenty British and is probably not nearly related. It is likely species are Holarctic and are fully described to have a female with simple tarsi and two- by him, while many others are discussed segmented cerci. in connection with related American species. Although Laffoon figured the ovipositor of only a few species, Laxtovka & Kidd (op. Mycetophila lubornirskii Dziedzicki cit.) used its structure in the ruficollis Group and there are good specific characters in the Mycetophila lubo~nirskii Dziedzicki, 1884: species covered here. Although several distinct 7, Table VIII, Figs. 19-22. species groups are discussed below, they all A male collected in Norfolk had distinct belong to Laffoon's Group F (both v and genitalia agreeing with Dziedzicki's (1884) a-d bristles on mid tibia, no a-d on hind figures of lubomirskii, described from 3 d tibia and at least apical third of second row collected in June in Bjelo-Russia. It is of anterior setulae on hind tibia dark brown) apparently scarce as the only more recent which is the largest, including about 20 British record was by Lundstrom (1 9 12) who found species. I d in Finland. The basistyle (ventral stylomere) has a dorsolateral lobe bearing a single long curved terminal spine as in the lunata Group, but The Mycetophila pictula Meigen Group lacks lateral spines on this lobe. It is nearest to the Nearctic clavata van Duzee but in This Group is characterized by a distinct Laffoon's (1957) key it runs most readily pre-apical wing spot just touching the tip of to napaea (Laffoon) as it has longish posterior vein R1, the male genital structure and in hairs on the hind coxae (separating it from the the female thickened fore tarsi and single- pictula Group) and largely dark brown ventral segmented cerci. All other known British setulae on the hind tibiae (excluding it from Mycetophila females have two-segmented the lunata/pectita Group). Both clavata cerci although some other Nearctic species and napaea belong to the Group which have a single segment. Thickened female fore includes the British species M.luctuosa Meigen tarsi are more frequent, also found in the which, however, lacks the dorsolateral lobe following: pumila Winnertz, unicolor of the basistyle of lubomirskii and clavata. Stannius, rudis Winnertz, strigata Staeger, In Landrock's (1927) key the British sigillata Dziedzicki, cingulum Meigen, specimen runs to nigrofusca Dziedzicki be- occultans Lundstrom and (only slightly) cause it has only two (instead of three) edwardsi Lundstrom. One British species, ventral bristles on the mid tibia and an bialorussica Dziedzicki, has thickened tarsi entirely dark abdomen but these characters in both sexes and agrees with the pictula may be variable, nigrofusca again being Group in the position of the pre-apical wing distinct on hypopygial structure. M.lubo- spot but has two-segmented female cerci. mirskii differs from pictula in the pre-apical It is more nearly related to strigatoides wing band being distinct only to the middle Landrock (=venusta Laffoon), a Holarctic of cell R5, although it is continued faintly species not recorded in Britain. beyond. The terminology used here in des- Edwards (1925b) recorded two species of criptions of Mycetophila species is based on this Group from Britain, i.e. pictula (as that of Laffoon (op. cit.). Studies of some British Mycetophilidae 85

Male. Head dark, palpi and basal antennal 2 strong v. Hind tibia with 4 strong d, inter- joints yellow; antennal flagellum dark but spersed with about 5 short weak bristles; basal 3-4 segments yellowish beneath. 0 a-d, 6-7 a, 3-4 short weak yellow p near Mesoscutum with three nearly fused dully tip. Tibial setulae all brown. Hind coxae with shining black stripes; yellow humeral areas, distinct posterior setae from basal third, narrow front margin and prescutellar spot longer apically, more than half length of extended as a central band on scutellum to external preapical bristle. base of metanotum. Scutellum and metano- Material examined. NORFOLK: Felbrigg tum (mediotergite) otherwise dark brown; Great Woods, 24 .X.1975, d (Zsmay). pleura dark brown except yellow proepister- num. Halteres yellow. Abdomen dark brown with small brownish yellow hypopygium Mycetophila autumnalis Lundst rom (Figs. 47-49). Legs yellow except for narrowly darkened tip to hind femora. Mycetophila autumnalis Lundstrom, 1909: Wings with median spot covering bases 60, Figs. 140-142. of cells R1, R5 and M1; preapical band just The recognition of lubomirskii led to the , . touching tip of vein R1 , filling end of cell R1, discovery of this species also introduced as then contracted basad, distinct to middle of British on a single male, apparently the first cell R5, faintly extended across m-fork, since the unique Finnish Type. Landrock 1 barely exceeding M2. Vein R with about ( 1927) briefly described autumnalis but 13 setulae below; M before r-m bare. Petiole omitted it from his key, in which the British of m-fork slightly shorter than r-m. Wing example runs to lubomirskii or bimaculata about 3.2 mm. dependent on whether the thorax is con- Mid tibia with 5 d, 1 a-d, 3 a, 4 weak p, sidered shining. The British male conforms to

FIGS. 44-49. Male hypopygia of Mycetophila species: autumnalis Lundstrijm, (44) ventral view; (45) external view of ventral stylomere; (46) external new of dorsal stylomere; lubomirskii Dziedzicki; (47) ventral view; (48) external new of ventral stylomere; (49) external new of dorsal stylomere. 86 Peter J. Chandler

Lundstrom's genitalia figures and to his Mycetophila bimaculata (Fabricius); Edwards, description except in having the thoracic 1925b: 639. stripes fused; the type had three closely Fungivora binlaculata (Fabricius); Laffoon, approximated stripes on a reddish yellow 1957: 277. ground. This character, however, varies in My cetophila pictula Meigen; Laffoon, 1965: other species (see below) and Edwards (1 928) 213. recorded pictula with well separated thoracic Ovipositor, Fig. 50. Examination of females stripes from Corsica. of this Group has confirmed their close re- The three-lobed basistyle distinguishes lationship. The male of pictula has a simple autumnalis from allied European species but triangular ventral lobe to its basistyle (figured similar structure is found in the Nearctic by Laffoon, 1957) and a weak dorsal lobe alata Guthrie. The proportions are different similar to tridentata but lacking the tridentate and alata also has a few setulae on the portion margin. Edwards (1925b), who reared it from of vein M before r-m. Poria vaporaria Cooke, said 'fairly common', Male. Head dark. Antennae brownish but it is unaccountably scarce in recent yellow on basal joints and base of first material. I have seen 18d and 21 9 mostly flagellar segment, rest of flagellum dark. from southern England, extending north to Thorax slightly shining, most of disc covered Inverness, but only l d and 39 collected by three fused pruinose blackish stripes; since 1950. broad brownish yellow humeral area connected with like coloured side margins. Scutellum dark except at basal corners. Metanotum and pleura nearly entirely Mycetophila forcipata Lundstrom blackish. Abdomen dark brown; hypopygium Mycetophila forcipata Lundstrom, 1913: 318, small, brownish yellow (Figs. 44-46). Legs Table XVI, Figs. 28 and 29. yellow but slightly darkened at tip of hind Mycetophila luteicauda Edwards, 19 13: 379, femora. Figs. 77 and 78. Wings with brown central spot covering Mycetophila forcipata Lundstrom; Edwards, Rs and bases of cells R5 and M1; pre-apical 1925b: 639. band just touching tip of vein RI, filling end Ovipositor, Fig. 5 1. Elongate cerci are found of cell RI, sharply contracted basally to a in both sexes. This is a frequent species which narrow band in cell R5, then at middle of develops in Piptoporus betulinus Fries; 208 cell deflected distally as a fainter shade and 15 9 have been seen from many parts of broadening across fork veins to faintly reach Britain and from Ireland. wing margin in cell Cu1 . Vein R with 13-1 5 setulae below; M before r-m with only 0--1 below at junction with r-m. Petiole of m-fork Mycetophila tridentata Lundstrom nearly equal to r-m. Wing length 3.5 mm. Mid tibia with 5-6 d (stronger apically), Mycetophila tridentata Lundstrom, 191 1, l a-d, 3 a, 3 p, 3 v (basal short). Hind tibia 415, Table XIV, Figs. 1 1 and 12. with 4 strong d, interspersed with 6-8 short Mycetophila tridentata Lundstrom; Edwards, weak bristles; 0 a-d, 8 a, 2-3 very weak 1926: 23 ; Edwards, 1941 : 80. yellow p near tip. All tibial setulae brown. This has a largely dull yellowish brown more Hind coxae practically bare posteriorly. humped thorax with three abbreviated brown Material examined. WEST SUSSEX: Dur- stripes. The previously unknown female is ford Heath, 26 .vii. 1974, d (Chandler). very similar to the male, differing only in the somewhat thickened joints 2-3 of the fore tarsi (less than in pictula and forcipata). Male genitalia, Figs. 58-60; ovipositor, Fig. My cetophila pictula Meige n 52. Sciara bimaculata Fabricius, 1805: 59 Although only the three specimens re- (preocc. Meigen, 1804: 92). corded by Edwards (1926, 1941) are in Mycetophila pictula Meigen, 1830: 299. BMNH, I have found tridentata widespread Studies of some British Mycetophilidae 87

FIGS. 50-57. Ovipositors of Mycetophila species in lateral view: (50) pictula Meigen; (51) forcipata Lundstrom; (52) tridentata Lundstrom; (53) bialorussica Dziedzicki; (54) lunata Meigen; (55) dziedzickii n0m.n.; (56) unipunctata Zetterstedt; (57) confusa Dziedzicki. in the south and have reared it from.Gano- The mesoscutum is black but for the yellow derma applanatum Karst., collected humeral areas as in pictula and forcipata, 14.ix.1964, dQ emerged 18.ix.1964, but is more strongly shining. Ovipositor, Farningham Wood, KENT. In all 14d and Fig. 53. It may have a northernlwestern 3 9 have been examined. New records are as distribution but only 4 have been seen. New follows: MIDDLESEX: Perivale Woods, records are: CORNWALL: Lelant, 2 .vi. 1974, d; BERKS: Windsor Forest, 28 .viii. 1912 (Yerbury, BMNH); ROSS: Ding- 7.vi.1973, Q, 20.vi.1974, d, 3.viii.1974, wall, l l .viii. 1909, d (King, BMNH); d, 2 .xi. 1975, d; WILTS: Savernake Forest, PERTHSHIRE: Foss Bridge, 22 .vi. 1974, Q 28. vi. 1971, 3 d; SUFFOLK: Brandeston, (Chandler). 1O.ix. 1972, d; BRECON: Llanddew, 26 .vi. 1976, d (all Chandler); HANTS: Mark Ash, 2.vi.1975, Q (Stubbs); CAMBS: Mycetophila confusa Dziedzicki Chippenham Fen, l .X.1946, d (Collin, HD). Mycetophila confusa Dziedzicki, 1884, 9, Table VI, Figs. 19--21 (captioned in- correctly as M.stolida Walker, corrected by Mycetophila bialorussica Dziedzicki Dziedzicki, 1886). Mycetophila bialorussica Dziedzicki, 1884, Mycetophila affluctata Edwards, 1941: 79, 14, Table VlI, Figs. 1-4. Figs. 9a-b, Syn.nov. 88 Peter J. Chandler

Laffoon (1957) drew attention to the close (not a female as stated by Becker, 1908) resemblance between genitalia figures of and found it to be conspecific with the confusa, briefly described by Dziedzicki British and European species punlila Winnertz from a single Bjelo-Russian male (compared (this synonymy to be established in an with stolida Walker) and affluctata, which account of the Canarian Mycetophilidae Edwards described from a British ~naleand under preparation). female (compared with unipunctata Meigen). The additional material of confusa differs Examination of this and other British from Edwards' type of affluctata in that the material has convinced me that these names mesoscutal stripes are not completely fused. apply to the same species. Also unipunctata sometimes has these stripes M.confusa is close to the Nearctic linlata blacker and more approximated than usual so Laffoon and to the common Holarctic uni- this character used by Edwards does not hold punctata Meigen. Laffoon stated, following good. Laffoon's character for limata (first published figures, that confusa and affluctata row of posterior setulae ventral to dorsal differed from limata in having two short bristles of hind tibia darker brown than stout setae on the posterior border of the succeeding rows; all setulae pale in urli- dististyle directly behind its point of attach- ~unctata)also applies to corlfusa but is not a inent. While only two are apparent from the very clear distinction. The male genitalia are angle figured, however, there is really a lobe distinct (figured by Edwards, 1941), i.e. bearing a row of about six close-set spines best the three strong spines on the basistyle of seen from a posterior view; Fig. 62 shows this unipunctata are represented by fine bristles aspect. in corlfusa and lirnata. The females are less Edwards (op. cit.) thought affluctata easily recognized but their ovipositors are was close to the Canarian fluctata Becker distinct (Figs. 56 and 57). There may be a (hence his choice of name) but he had not constant difference in the antennae (segments seen fluctata, which has not been figured. I a little more than twice as long as broad in have examined Becker's type, which is a male unipunctata, somewhat less in confusa).

59 60 6 2 FIGS. 58-62. Male hypopygia of Mycetophila species: tridentata Lundstrom, (58) ventral view; (59) internal view of right stylomeres; (60) postero-external view of right stylomeres; confusa Dziedzicki, (61) ventral view; (62) postero-internal view of right stylomeres. Studies of some British Mycetophilidae 89

Material exanlined. Holotype d of Myceto- Meigen Group such as alea Laffoon (=guttutu phila affluctata Edwards, SUFFOLK: Eris- Dziedzicki of Edwards) and signatoides well, 24 .X. 1934 (Collin, BMNH). SUFFOLK: Dziedzicki, common species with more or less Brandeston, 3.X. 1972, d (Chandler); separated black stripes on a strongly shining CAMBS.: Chippenham Fen, 16.viii. 1940, 9 yellow mesoscutum. They developin terrestrial 21.ix.1941, 4d, 29; 24.ix.1941, 2d, 39; gill fungi and the rearing records of lunata 13.X, 1943, 6 (Collin, HD). from Agarics no doubt refer to them or other related species. Walker (1836, 1856) recorded lunata from the British Isles but at least The Mycetophila lunata Meigen Group Haliday's Irish material was alea (Chandler, 1976). Lundstrom (1 906) and possibly some This corresponds to the Nearctic pectita earlier Scandinavian authors used the name Johannsen Group, said by Laffoon (1957) lunata for finlandica Edwards. to include three Nearctic species (pectita The present usage of lunata follows Johannsen, subita (Laffoon) and contigua Winnertz (1863) according to Dziedzicki's figures, an interpretation followed by most d. Walker) and the European obscura Dzied- zicki (a homonym renamed below); he did not later authors. Meigen (1804) said it was mention lunata heigen although its close common on ivy blossom and developed in , relationship can be seen from Dziedzicki's bark encrusting fungi, which would fit the figures. Edwards (1 913) said that obscura species dealt with here. Edwards (1924) might be a dark form of lunata because of did not mention lunata in his review of similarity in hypopygial structure, while Meigen's types but SCguy (1 940) described a Laffoon refers to pectita and obscura as an female specimen said to be the type of lunata allopatric species pair. M.lunata, however, and there is nothing in his description to rather than obscura is nearer to pectita to conflict with this identification. It is not which it runs in Laffoon's key. I have not possible to say how many records can be considered lunata and pectita conspecific, accepted but the true lunata sensu Dzied- although this is possible, because of small zicki is probably widespread but local in differences in the shape of the dorsal western and central Europe. processes of the dististyle. I became aware of lunata as a British Laffoon stated that he had examined species when Mr R. E. Evans sent to me reared European material of obscura but did not material which ran in Edwards' (1925b) key mention that it differs from the three Nearctic to firzlandica because of their separate species in possessing a row of setulae below thoracic stripes and lunata is coupled with vein M before r-m, also completely absent in firzlandica in Landrock's (1927) key, but the lunata. Because of this character, obscura genital structure is quite different. I later runs in Laffoon's key to strigata Staeger, found, however, that lunata was mixed with which is not closely allied. Otherwise, obscura obscura in collections and some, especially closely resembles contigua, differing again that from Norfolk, had fused stripes with most obviously in the shape of the dorsal only the shoulders yellow and the thorax processes of the dististyle. more shining; these run to obscura in Edwards' key. The genitalia of both have a spinose dorsolateral lobe to the basistyle, also found in the Nearctic species of the Mycetophila lunata Meigen group. M.lunata has the apical spine of this Mycetophila lunata Meigen, 1804: 90; lobe markedly curved and 7-8 lateral spines Winnertz, 1863: 931; Dziedzicki, 1915, as in pectita; obscura has the apical spine Plate XVIII, Figs. 283-286 (based on nearly straight and only 4-5 lateral spines. Winnertz' collection). The ventral lobe is also distinctly longer in It is necessary to discuss the interpretation of obscura while the lobes are about equal in this name. Many nineteenth century authors lunata. There is a greater difference in the cited lunata but most probably refer to lunata structure of the delicate dististyles. of Fabricius (1805), i.e. species of the signata Male and female. Head dark. Palpi and 90 Peter J. Chandler basal antenna1 joints, including whole of first not nearly reaching back to vein RI, arching flagellar segment yellow, rest of flagellum basad in cell R5, then extending faintly dark. Mesoscutum with three dully shining across median fork, ending just beyond vein black stripes, narrowly separated or com- M2. Both wing markings much stronger than pletely fused; median stripe reaches fore in nbscura. Vein R with 9-1 1 setulae below; margin but broad humeral area and adjoining vein M before r-m bare. Petiole of m-fork side margins of mesoscutum yellow. A yellow short, less than half length of r-m. prescutellar spot (larger when the stripes are Mid tibia with 6 d, 1 a-d, 3 a, 2-3 strong separate) extended as a median band onto p (2 short weak yellowish bristles above scutellurn. Rest of scutellum, metanotum and them), 3 v. Hind tibia with 4 strong d (inter- most of pleura dark; pronotum and pro- spersed with about 6-8 short weak bristles), episternum yellow. Abdomen dark with 0 a-d, 7 a, 4-8 short yellow p near tip. narrow yellow hind margins to segments; Mid tibia with anterior setulae brown, hypopygium (Figs. 63-66) and ovipositor posterior yellowish. Hind tibia with first 1-2 (Fig. 54) brownish yellow. Legs pale yellow rows mainly brown, rest of anterior rows only with tips of mid and hind femora narrowly brown on apical quarter to third, ventral dark brown. brown only near tip and posterior all yellow. Wings with large dark brown median spot Hind coxae with several long posterior hairs in bases of cells R1, R5 and M1 ; a narrow near tip, nearly as long as external preapical, prcapical band filling the end of cell R1 but only shorter hairs basally. Female fore tarsi

FIGS. 63-69. Male hypopygia of Mycetophila species: lunata Meigen, (63) ventral view; (64) external view of dorsal lobe of left ventral stylomere; (65) internal view of ventral lobe of left ventral stylomere: (66) internal view of left dorsal stylomere: dziedzickii nom.n., (67) ventral view; (68) external view of left ventral stylomere; (69) external view of left dorsal stylomere. Studies 01'some British Mycetophilidae 9 1

simple (slightly thickened on segments 2-3 and base of antennae (including less than basal in pectita). half of first flagellar segment) yellow. Meso- Wing length, d 3.5-4.0 mm; ? 4.1- scutum with three fused shining brown 4.3 mm. stripes, leaving a large yellow humeral area not Material examined. NORFOLK: Wheatfen connected with small yellow postalar and Broad, 31 .X.1975, d (Irwin), 30.iv. 1976, d prescutellar spots, the latter joining median (Ismay); SUFFOLK: Boyton, 27 .viii. 1907, yellow scutellar band. Metanotuni and most d (Collin, HD); CAMBS.: Chippenham Fen, of pleura dark brown but pronotum, -pro- ll.viii.1940, d, 24.viii.1940, d, 21 .ix.1941, episternuni and upper part of katepisternum d, 24.ix.1941, d (Collin, HD); HUNTS.; yellow. Abdomen dark brown with narrow Monk's Wood, 29 .ix ,1972, d (Cole); NOTTS.: yellow hind margins to segments; hypopygium Strelley, 21 .ix. 1922, d (Edwards, BMNH); (Figs. 67-69) and ovipositor (Fig. 55) WARWICKS.: Dumble Wood, reared ii. 1975 brownish yellow. Legs yellow; tips of mid and ex Coniophora puteana Karst., l d, 39 hind femora narrowly brown. (Evans). Wings with small brown median spot concentrated in base of cell R5 but extending over Rs and into upper part of cell M1; preapical band fainter, filling tip of cell R1 Mycetopllila dziedzickii n0rn.n. and nearly reaching back to vein R1 I t (approaching pictula Group in this respect), Mycetophila obscura Dziedzicki, 1884: 8, arched basad in cell R5 but becoming very Table VIII, Figs. 30-33, nec Walker, faint beyond middle of cell, disappearing 1848: 101. just beyond vein M2. Vein R with 9-12 This new name is proposed because of the setulae below; M before r-m with about homonymy stated above, which was first 7 setulae on apical half below. Petiole of mentioned by Laffoon (1957: 282). m-fork short, half or slightly less length of M.obscura Walker is a Phror~iaspecies which r-m. cannot be identified because of the poor Mid tibia with 5-6 d, 1 a--d, 3 a, 2-3 v condition of the Type (in BMNH). (basal when present weaker), 3-4 p (1-2 Since Dziedzicki figured the hypopygium upper very short and weak); anterior setulae when describing obscura from a single Polish brown, posterior yellow. Hind tibia with male, most published records are probably 4-5 long interspersed with 3-5 short d, correct although some evidently apply to 0 a-d, 8 strong a (sometimes one short lunata. It is widespread in Europe but has not additional basal to them), 2-3 very short been reared. Edwards (19 13, 1925b) quoted p near tip; first anterior row of setulae brown, several British localities; his Baldock and rest only brown on apical third to half, Shefford specimens can be confirmed but the posterior setulae all yellow. Hind coxae with Strelley example was lunata (see above). many pale posterior hairs becoming longer on The ovipositors of lunata and dziedzickii apical half. Female fore tarsi simple. (not previously figured) agree with the Wing length, d 3.0-3.5mm; ? 3.7- Nearctic species of the group in the possession 4.0 mm. of produced terminal lobes to tergite IX, Material examined. HANTS.: Leckford, covering part of the basal joint of the cerci, d? in copula at ivy flowers, 5.x. 1975 differing in this respect from other Myceto- (Chandler); I. of WIGHT: Freshwater, d? at phila species. The shape of this lobe and of windows, 5 .ix. 1948 (K. G. Blair); HERTS.: the cerci are closely similar (according to Baldock, v. 1916, d (Edwards, BMNH); Laffoon's figures) in lunata and pectita and BEDS.: Shefford, viii. 1913, l d, 2 9 there is a similar resemblance between dzied- (Edwards, BMNH); SUFFOLK: Newmarket, zickii and contigua, no obvious differences Sussex Lodge, 12.ix.1888, ? (Verrall, HD), being apparent in either case. This tends to 17 .ix. 1922, d (Collin, HD); Tuddenham, confirm the view that two very closely related 16.ix.1891, d (Verrall, HD); Hollesley, allopatric species pairs are involved. 17 .ix. 1909, d (Yerbury, HD); INVERNESS: Male and female. Head dark with palpi Aviemore, 20.vi. 1903, d (King, BMNH). 92 Peter 1. Chandler

Acknowledgments Dziedzicki, H. (1923) Revue des espices europ6en- nes du genre Anatella Winnertz avec la description des deux genres nouveaux, Heteropygiurn I am indebted to the authorities of the British et Allophallus. Discipl. biol. Arch. Soc. Sri. Museum (Natural History) and the Hope Varsoviensis, 1, (1922), 1-8, 2 pl. Department of Ento~nology, Oxford Uni- Edwards, F.W. (1913) Notes on British Myceto- versity Museurn, for the opportunity to philidae. Trans. ent. Soc. Lond. 1913, 334-382. examine material housed by them and for Edwards, P.W. (1921) Diptera Nematocera from Arran and Loch Etive. Scot. Nut. 1921, 59-6 1, loan of selected specimens. The following 89-92, 121-125. collectors have also kindly loaned to me many Edwards, F.W. (1924) Notes on Meigen's fungus batches of Mycetophilidae collected by them- gnat types. Encycl. Ent. (B2) Dipt. 1, 13-17. Mr J. H. Cole, Mr R. E. Evans, Mr J. W. Ismay, Edwards, F.W. (1925a) Notes on the types of Mycetophilidae (Diptera) described by Staeger Mr A. G. Irwin, Mr G. Hosie and Mr A. E. and Zetterstedt. Ent. Tidskr. 45, 160-168. Stubbs. They have generously allowed me to Edwards, F.W. (1925b) British fungus gnats (Dip- bring together the results of their efforts tera, Mycetophilidae) with a revised generic and to make full use of data obtained by classification of the family. Trans. ent. Soc. them. Dr Matile of the MusCum National Lond. 1924, 505-662. L. Edwards, F.W. (1926) The Wood collection of dlHistoire Naturelle, Paris, has (in litt.) British Diptera and Lepidoptera. Entomologist's discussed and advised on some taxonomic mon. Mug. v aspects. Edwards, F.W. (1928) The nematocerous Diptera of Corsica. Enc~~cl.Ent. (B2) Dipt. 4, 157-189. Edwards, F.W. (1933) Exechia bicincta Staeger (Mycetophilidae), a Trichocera (Trichoceridae) and Tipula vittata Meigen (Tipulidae) with References abnormal characters (Dipt.). J. ent. Soc. S. Engl. 1, 55-56. Becker, T. (1908) Dipteren der Kanarischen Inseln. Edwards, F.W. (1941) Notes on British fungus Berlin Mitt. Zool. Mus. 1908, 1-80. gnats (Dipt., Mycetophilidae). Entomologist's Bukowski, W. (1949) New species of fungus gnats mon. Mug. 77, 2132,6782. (Diptera, Fungivoridae) from the Crimea (in E'abricius, J.C. (1805) Systema antliatorum secun- Russian). Ent. Oborr. 30,405 -409. dum ordines, genera, species, adjectis synonymis, Burghele-Balacesco, A. (1966) Les Mycetophilidae locis, observationibus, descriptionibus. Bruns- (Diptires) cavernicoles de la collection Bio- vigae, Reichard. speologica (IVe-VllIe series des "Grottes Johannsen, O.A. (1909) Diptera. Family Myceto- visittes." Int. J. Speleol. 2, 3 19-334. philidae. Pasc. 93, 141 pp. in Wytsman, P. (Ed.) Burghele-Balacesco, A. (1972) Contribution a Genera Insectorum, Brussels. l'itude des Mycetophilidae des grottes de Johannsen, O.A. (1912) The fungus gnats of North Roumanie avec la description de deux espices America. Part 1V. 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(1884) Przyczynek do fauny owadow boheniica Lactovka and Dynatosoma nigro- dwuskrzydtych. Gatunki rodzajow: Mycothera, maculatum Lundst. new to Britain and notes on Myce tophila, Staegeria. Pamie tnika Firyjograficr- other little known fungus-gnats (Dipt., Myceto- nego, 4, 298-324. German translation without philidae). Entomologist, 103, 10-17. Plates: Beitrag zur Fauna der zweiflugeligen Lackschewitz, P. (1937) Die Fungivoriden des Insecten (Arten der Gattungen Mycothera, Ostbaltischen Gebietes. Arb. Naturforsch. Ver. Mycetophila und Staegeria) (1886) Wien ent. Riga, 21, 1-47. Zeit. 5, 153-156, 189-190, 229-231, 251- Laffoon, J. L. (1957) A revision of the Nearctic 253,265-266,326-327, 346-347; 6, 37-43). species of Fungivora (Diptera, Mycetophilidae). Dziedzicki, H. (191 5) Atlas des organes ginitaux IowaState Coll. J. Sci. 31, 141-340. des types de Winnertz et des genres de sa collec- Laffoon, J.L. (1965) Family Mycetophilidae, pp. tion de Mycitophiles. Pub. Soc. Sci. 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