Revision of Holarctic of Meigen 195 rounded (but shrivelled in some); aedeagus (as ater was based on material from Megerle. in Fig. 10A) with lateral prongs, directed antero- Megerle's material was destroyed in the fire of laterally, distal portion a simple, gradually 1848 at the Vienna Museum (Pont, 1986). We tapering projection, about twice as long to examined three specimens from the Winthem markedly longer than lateral prongs, very apex collection (in NHMW) which were labelled as directed ventrally. 'ater', but unfortunately not in Meigen's hand- Female adult. Descriptive statistics: see writing. These could therefore not be considered Tables 8—13. Similar to male except for usual as material and the type material of S.atra sex differences and as follows: is considered to be lost. Head: mandible serrate. The description of both S. micronyx and Legs: coloration pattern as indicated in Fig. S. albitarsis by Kieffer (1919) as having females 5D; strong bristles of femora, tibiae distributed with a brilliantly shiny thorax and short, equal as follows: present or absent ventrally on fore, hind claws can only refer to S.atra as described mid femora, ventrally on hind femur; claws of here. The neotypes are proposed to ensure hind leg equal, small. stability of the names applied to Palaearctic Wing: macrotrichia restricted to apical species. margin. It is clear from Kieffer's (1919) list of nine Genitalia (as in Fig. 12D): sternite 9 truncate localities for S.spinosipes that he based his de- medially to somewhat truncate but with antero- scription on a number of specimens. The only medial margin developed, pointed; two available material of Kieffer's type series spermathecae, additional spermathecal duct was the lectotype and a paralectotype housed terminating in pigmented apex. in the BMNH. The single paralectotype of Pupa. Described under generic description. S.spinosipes, however, is a female adult of Distribution and bionomics. S.atra is known S. morio. from England to Latvia, south to Belgium and Kieffer's (1919) description of S.spinosipes Hungary (Fig. 18B). Adults have been collected does not quite fit the description of S.atra here from 13 May to 6 July. in that the femora and tibiae of S.atra are nearly Little can be said about the bionomics of this completely darkly pigmented. Kieffer (1919) species. Goetghebuer (1936b) noted that adult reported S.spinosipes with the fore and mid S.atra were associated with rivers in northern femora and tibiae as mostly yellow. However, Belgium but we were unable to confirm his the lectotype here designated fits the description identifications. Two specimens from Britain had of S.atra in this study. labels which noted that the specimens came One female pupa (ZSMC, material from from fens (Chippenham, Wood Walton Strenzke, 1950) we examined contained a (Hunts)). Strenzke (1950) collected a pupa from pharate adult and was somewhat dirty. What the shore of Schohsee, F.R.G. few differences between this specimen and other Taxonomic discussion. The association of the available material of Serromyia pupae we could males and females was based on specimens detect (size and number of spiracular openings) collected at the same place and time in Latvia are noted in the generic description. The label (by Remm), Belgium (by Goetghebuer: types on the slide noted 'femorata (?) morio?' and of S.nitens) and Amsterdam (by Meijere, 1946: '246' (therefore collected at Schohsees accord- reported as S.atra). ing to Strenzke's (1950) records). Neotypes have been designated for Cera- The male genitalia of S.atra are indistinguish- topogon ater, Serromyia micronyx and able from those of S. morio. However, the Serromyia albitarsis because type specimens characters given in the keys and significant dif- could not be located and have probably been ferences in the ratio of antennal flagellomeres destroyed. 10/11 indicates that these are separate species. The description of ater by Material examined. 59 males, 11 females and Meigen (1818) and the accompanying plate 1 female pupa. drawn by Meigen (in Morge, 1975) showing the Derivation of specific epithet. The name ater extent of leg pigmentation, corresponds to the (black) presumably refers to the overall dark concept of S.atra as proposed where. Meigen appearance of adults of this species noted by (1818) noted that his description of Ceratopogon Meigen. 196 A. Borkent and B. Bissett

Serromyia bicolor Borkent sp.n. the uniquely (amongst Holarctic Serromyia) bicoloured palpus. Diagnosis. Male and female: only Palaearctic The male of S. bicolor is unusual in a number species with palpal segments 1-3 pale, segment of characters and is the most distinctive of all 4 light brown and segment 5 dark brown. the Holarctic species (other than mangrovi). Description. Male adult. Descriptive statistics: Colour pattern of both male and female palpus, see Tables 2-7. the peculiar apex of the aedeagus, the sword- Head: dark brown; antennal flagellomere 10 like parameres and a markedly slender (for a with plume arranged in more than one whorl; species of Serromyia in the Holarctic) hind palpus with segments 1—3 pale, segment 4 light femur of the male are all unique to this species. brown, segment 5 dark brown. Havelka & Caspers (1981) partially described Thorax: dark brown; scutum pruinose. this species under the name S. subinermis Kieffer Legs: coloration pattern as indicated in Fig. (males are those of S.bicolor as recognized here, 3E; strong bristles of femora, tibiae distributed females are actually those of S. bicolor and as follows: ventrally on mid femur, ventrally S.subinermis). However, the name subinermis on hind femur, dorsally on hind tibia; ventral cannot apply to the present species as Kieffer bristles on hind femur arising from slightly (1919) noted that S. subinermis had spines developed tubercles; hind Taj straight; Ta4 setae (strong bristles) on the fore and mid femora in with slight curve or some slightly sinuate near the female and none in the male. Male S.bicolor their apex. have strong bristles on the mid femur only Wing: slightly infuscated with veins of cells and in the female these were either present or r1; r2+3 darkly pigmented. absent. As far as we can discern, there are no Abdomen: dark brown. Palaearctic names which could apply to this Genitalia (Fig. 9C): gonostylus sinuous at species and have accordingly described it as midlength, tapering gradually for basal half, new. with swollen apex; paramere (Fig. 9B) blade- In their drawing of the male genitalia of like, with pointed apex, directed posteriorly; S.bicolor (as S.subinermis), Havelka & Caspers aedeagus (Fig. 9A) lacking lateral prongs, distal (1981) misinterpreted the apex of the aedeagus portion truncate, with jagged margin, with as elongate and slender. posterolateral lobes, extreme apex directed Types. Holotype, 8 adult on microscope posteriorly. slide, labelled 'Holotype Serromyia bicolor Female adult. Descriptive statistics: see Borkent, Bonn, Federal Republic of , Tables 8-13. Similar to male except for usual June 7, 1976, P. Havelka, A551' (PEHC); sex differences and as follows: allotype, 9 adult on microscope slide, labelled Head: mandible serrate. as for holotype but collected 24 May 1976 Legs: coloration pattern as indicated in (PEHC); paratypes: 28 from holotype locality, Fig. 5B; strong bristles of femora, tibiae dis- dated 10.v. 1976 (CNCI), 11.v. 1976, (PEHC); tributed as follows: present or absent ventrally 19 U.S.S.R., Caucasus, Dilizan, 5.vi.l969 on mid femur, ventrally on hind femur; claw of (CNCI). hind leg single, elongate, with basal tooth. Derivation of specific epithet. The name Wing: macrotrichia restricted to anteroapical bicolor (two-coloured) refers to the distinctive margin. bicoloured palpus of males and females of this Genitalia (as in Fig. 12D): sternite 9 truncate species. medially, with anteromedial margin developed, pointed; two spermathecae, additional sperma- thecal duct terminating in pigmented apex. (Meigen) Distribution and bionomics. S.bicolor is known from Bonn, West Germany, and from Ceratopogon femoratus Meigen 1804: 28. Dilizan in the Caucasus Mountains of the Lectotype, here designated, 8 adult on Armanskaja S.S.R. (Fig. 18B). Adults have microscope slide, labelled 'Lectotype Cera- been collected from 10 May to 7 June. topogon femoratus 8 Meigen, ex coll. Meigen Taxonomic discussion. We have associated 291/40 (158a), prep. J. Clastrier, 1985 baume the single female with the males on the basis of phenole, Museum , Serromyia femorata Revision of Holarctic species of Serromyia Meigen 197

Det. A. Borkent' (MNHN); paralectotypes: Clastrier 1963: 61 (in part). 2 9 labelled as for holotype (MNHN). Staeger Ceratopogon morio: authors, not Fabricius. 1839: 598 (in part). Zetterstedt 1838: 822 (in Staeger 1839: 598 (in part). Zetterstedt 1850: part), 1850: 3665 (in part), Schiner 1864: 584 3666 (in part). (in part). Chironomus femoratus: Fabricius 1805: 45. Diagnosis. Male: only Palaearctic species with Prionomyia femorata: Stephens 1829: 237. three rows of strong bristles on the fore femur, Ceratolophus femoratus: Kieffer 1899: 69. scutum pruinose and a pale wing. Female: only Johannseniella femoratus: Williston 1907: 1. Palaearctic species with palpus darkly pig- Serromyia femorata: Kieffer 1901: 157. Kieffer mented, the scutum pruinose, nearly the entire 1906: 65. Goetghebuer 1934: 61 (in part). fore and mid tibiae yellow with only very apicies Edwards 1926: 410 (in part). pigmented in some, strong lateral bristles absent Ceratopogon armatus Meigen 1818: 83. Lecto- from hind coxa and a pale wing. type, here designated, 6 adult on microscope Description. Male adult. Descriptive statistics: slide, labelled 'Lectotype Ceratopogon see Tables 2—7. armatus 6 Meigen, ex coll. Meigen 291/40 Head: dark brown; antennal flagellomere 10 (161), prep. J. Clastrier, 1985 baume phenole, with plume arranged in more than one whorl; Serromyia femorata Det. A. Borkent, palpus dark brown. Museum Paris' (MNHN). Thorax: dark brown; scutum pruinose. Prionomyia armata: Stephens 1829: 238. Legs: coloration pattern as indicated in Figs Ceratolophus armatus: Kertesz 1902: 157. 3F, 4A; strong bristles of femora, tibiae distrib- Serromyia armata: Kieffer 1906: 65. uted as follows: anteriorly, ventrally, posteriorly Ceratopogon foersteri Meigen 1838: 21. Neo- on fore femur, dorsally, posteriorly on fore type, here designated, 8 adult on microscope tibia, present or absent anteriorly on fore tibia, slide, labelled 'Neotype Serromyia foersteri anteriorly, ventrally, posteriorly on mid femur, Meigen, 8, Newcastle-u-Lyme, Staff., posteriorly on mid tibia, present or absent an- England 3-VI-1960, J. R. Vockeroth CNC teriorly or dorsally on mid tibia, anteriorly, No. 20249, Serromyia femorata Det. A. ventrally on hind femur, dorsally on hind tibia; Borkent' (CNCI). ventral bristles on hind femur arising from Ceratolophus foersteri: Kertesz 1902: 157. slightly developed tubercles; hind Taj straight Serromyia foersteri: Kieffer 1906: 65. or with slight basal curvature; Ta4 setae straight Ceratopogon flavipes Gimmerthal 1847: 144, or with slight curve. not Ceratopogon flavipes Meigen 1804: 28. Wing: pale or slightly infuscated with veins of Neotype, here designated, 8 adult on micro- cells r1? r2+3 very lightly pigmented. scope slide, labelled 'Neotype Ceratopogon Abdomen: dark brown. flavipes Gimmerthal, 8, Oxford, England, 6- Genitalia (Fig. 9F): gonostylus with outer V-1953, J. R. Vockeroth, Neotype CNC No. margin evenly curved, tapering gradually 20272, Serromyia femorata, Det. A. Borkent' for basal half to tapering gradually to rounded (CNCI). New synonym. or somewhat swollen apex; paramere (Fig. Palpomyia flavicrus Kieffer 1906: 63. New name 9E) with apical half markedly swollen, apex for Ceratopogon flavipes Gimmerthal. New rounded; aedeagus (Fig. 9D) with lateral Synonym. prongs, directed posterolaterally, to dorso- Serromyia flavicrus: Remm 1988: 35. lateral^, distal portion a simple, slender pro- Serromyia inermipes Kieffer 1919: 73. Neo- jection, about twice to markedly longer than type, here designated, adult on microscope lateral prongs, extreme apex directed ventrally. slide, labelled 'Neotype Serromyia inermipes Female adult. Descriptive statistics: see Kieffer, Brockenhurst, Hants [= Hampshire], Tables 8—13. Similar to male except for usual England 3-V-1951, J. R. Vockeroth, CNC sex differences and as follows: No. 20273, Serromyia femorata Det. A. Head: mandible serrate. Borkent' (CNCI). Legs: coloration pattern as indicated in Figs Ceratopogon rufitarsis: authors, not Meigen. 5B, C; strong bristles of femora, tibiae distrib- Meigen 1818: 83 (in part). uted as follows: present or absent ventrally on Serromyia europaea: authors, not Clastrier. fore femur, present or absent ventrally on mid 198 A. Borkent and B. Bissett femur, present or absent at extreme apex of The specimens which Edwards (1920) noted mid tibia, ventrally on hind femur; claw of hind as preying on pulchripes Verr. and leg single, elongate, with basal tooth. Bezzia ornata (Meigen) were examined and are Wing: macrotrichia absent or a few restricted indeed members of S.femorata. to very apical margin. The following are literature records of prey Genitalia (as in Fig. 12D): sternite 9 trunc- or feeding for which we were unable to examine ate medially to somewhat truncate but with the original material and confirm the identifi- anteromedial margin developed, pointed; two cations. Edwards (1923) noted Trichocladius spermathecae, additional spermathecal duct sp. () as prey. Gad (1951) re- terminating in pigmented apex (one specimen ported S.femorata attacking lsohelea sociabilis with three fully developed spermathecae (Goetghebuer) and Culicoides impunctatus present). Goetghebuer. Goetghebuer (1949) reported Distribution and bionomics. S.femorata is that S. femorata males fed on flowers and females known from northern Fennoscandia south to preyed on Metriocnemus (Chironomidae). northern Spain, northern Yugoslavia and Edwards (1926) noted that male S.femorata Moscow (Fig. 19A). The locality in the Kola swarm at mid-day and Downes (1955) reported Peninsula shown on the map is based on a female swarms composed of 3—20 individuals specimen with no further specific distributional of S.femorata under the tips of alder branches data. Adults have been collected from 6 May to at Loch Lomond, Scotland. Although these 14 August. records are consistent with the distribution Havelka (1978, 1979) and Gil Collado (1957, of this species, we were unable to examine 1985) noted records of S.femorata (Meigen) any of this material and thereby confirm the from Spain. We have examined one female identifications. from Santander housed in the MZLU but have Taxonomic discussion. The type of Cera- been unable to confirm these previous records. topogon foersteri Meigen is not present in Paris Remm (1967) reported two specimens of (Clastrier, pers. comm.) or in Halle (Dorn, S.femorata from the Causcasus and these pers. comm.) and we have accordingly desig- identifications, too, could not be confirmed. nated a neotype for the species. Meigen's (1838) Strenzke (1950) noted the presence of larvae description is so general that this species might of S. femorata in lake-side mosses in Germany actually be one of any number of European but we have been unable to confirm these species. His drawing of the female (in Morge, identifications (material was examined, see 1975) shows a unique leg coloration with apically under generic description). darkened fore and mid femora and completely We examined five pairs of S.femorata col- pigmented fore and mid tibiae. However, some lected in copula from Norway, and pinned specimens of S.femorata may have Britain which allowed for confident association somewhat darkened tibiae (as in Fig. 5C). of the sexes. Previous workers (Goetghebuer, 1920, 1934; One of the mated pairs (from Sweden) had Remm, 1988) have considered this species to be the male abdomen broken between the sixth a synonym of S. femorata and the designation of and seventh abdominal segments with the the neotype here will establish this. male's genitalia firmly attached to that of the The type of Serromyia inermipes Kieffer could female. This indicates a mating behaviour not be located. Remm (1988) considered the similar to that recorded for various species of name a synonym of S.femorata although the Heteromyiini, Sphaeromiini and Palpomyiini general description given by Kieffer (1919) (Downes, 1978). Edwards (1920) noted that he could apply to several European species. We collected mating pairs of S.femorata in which have designated a neotype to confirm Remm's the females were feeding on males. We were synonymy. unable to locate the specimens upon which these Remm (1988) recently placed Ceratopogon observations were based and therefore cannot flavipes Gimmerthal as a species of Serromyia. confirm the identification. The presence of Based on the brief description given by the broken male abdomen and attached male Gimmerthal (1847), in which he noted that the genitalia noted above would correspond to the underside of the hind femora of the two female presence of such a feeding behaviour. syntypes were spinose, this is probably correct. Revision of Holarctic species of Serromyia Meigen 199

The rest of the description, however, is so gen- bristles on hind femur arising directly from flat eral that it does not allow for a confident place- cuticle or from very slightly developed tubercles; ment of the name. The mention that the fore hind Tai with marked basal bend; Ta4 setae and mid 'Schenkel' [probably = femur] and straight or with slight curve. 'Gelenke' [probably = base of tibia] were brown Wing: pale. is approximately a condition occurring in some Female adult. Descriptive statistics: see female S.femorata. The designation of the Tables 8-13. neotype here places Ceratopogon flavipes Legs: strong bristles of femora, tibiae dis- Gimmerthal as a synonym of S.femorata. tributed as follows: apically on mid tibia, Goetghebuer (1922a) noted that the syntype mventrally on hind femur. series of Ceratopogon femoratus was composed Genitalia: sternite 9 truncate medially. of one male and four females. We examined Distribution and bionomics. S. mangrovi is only the male and two females. known only from the Sinai Peninsula, Egypt. The male paratype of S.europaea from Nor- The specimens we examined were collected way and the male paralectotype of S. rufitarsis from 4/5 June to 18/19 November at a light were actually specimens of S.femorata. trap. Edwards (1926) suggested that female S.fe- Taxonomic discussion. Although Delecolle morata lack macrotrichiae on the wing mem- & Braverman (1987) designated only nine brane but this is true for only some specimens. males and nine females as paratypes, they also Material examined. 199 males and 246 examined an additional thirteen males and 122 females. females from the Sinai. Derivation of specific epithet. The name Material examined. Two males and two femoratus (pertaining to the femora) pre- females collected at E-Shira el Gharkana sumably refers to the swollen hind femur of and Ras Muhammad, Sinai Peninsula, Egypt members of this and several other genera of (deposited in CNCI). . Derivation of specific epithet. The name mangrovi refers to the mangrove swamp where Serromyia mangrovi Delecolle & the specimens were collected. Braverman Serromyia mangrovi Delecolle & Braverman 1987: 57. Holotype, 9 adult, not seen, from Serromyia morio (Fabricius) EGYPT: Sinai, Ras Muhammad (MZSF); Culex morio Fabricius 1775: 800. Neotype, here allotype, 8 adult, not seen, labelled as for designated, 8 adult on microscope slide, holotype (MZSF); paratypes (not seen): labelled 'Neotype Serromyia morio Fabricius, EGYPT: 9

Serromyia femorata: authors, not Meigen. Tables 8-13. Similar to male except for usual Edwards 1926: 410 (in part). Goetghebuer sex differences and as follows: 1934: 61 (in part). Head: mandible serrate. Serromyia atra: authors, not Meigen. Legs: coloration pattern as indicated in Figs Goetghebuer 1934: 61. 4E, 5E; strong bristles of femora, tibiae dis- Serromyia spinosipes: authors, not Kieffer. tributed as follows: absent or present ventrally Kieffer 1919: 72. on fore femur, absent or present on mid femur, Serromyia ledicola: authors, not Kieffer. Remm ventrally on hind femur; claw of hind leg single, 1969: 214 (in part). elongate, with basal tooth. Ceratopogon flavicornis: authors, not Staeger. Wing: macrotrichia with very few to many, Zetterstedt 1850: 3667. restricted to apical margin. Genitalia (as in Fig. 12D): sternite 9 trunc- Diagnosis. Male: only Palaearctic species with ate medially to somewhat truncate but with fore femur with three longitudinal rows of strong anteromedial margin developed, pointed; 2 bristles and with the prongs on the aedeagus spermathecae, additional spermathecal duct directed anterolaterally. Female: only Palae- terminating in pigmented apex (one specimen arctic species with the scutum lacking pruinosity, with 3 fully developed spermathecae). the mid femur distinctly paler basally, and with Distribution and bionomics. S.morio is known an elongate, single hind claw. from England to Estonia south to , Description. Male adult. Descriptive statistics: Austria and Greece (Fig. 19B). Adults have see Tables 2—7. been collected from 7 May to 16 August. The Head: dark brown; antennal flagellomere 10 extension into August was due to two speci- with plume arranged in more than one whorl; mens, one female from Hiiumaa Is., Estonia, 8 palpus dark brown. August and a male from , Federal Thorax: dark brown; scutum bare of Republic of Germany, 16 August. Otherwise pruinosity. the latest date of collection is 18 July. It is Legs: coloration pattern as indicated in Fig. possible that the two August specimens were 4A; strong bristles of femora, tibiae distributed mislabelled. as follows: anteriorly, ventrally, posteriorly on The most southerly locality of S. morio is fore femur, dorsally, posteriorly on fore tibia, Lake Ochrid, Macedonia, Yugoslavia. Edwards present or absent anteriorly on fore tibia, (1928) recorded this species from Calvi, Corsica, anteriorly, ventrally, posteriorly on mid femur, but the single female was actually a specimen of posteriorly on mid tibia, present or absent S. subinermis. anteriorly, dorsally on mid tibia, anteriorly, Remm (1967) reported S.morio from the ventrally on hind femur, dorsally on hind tibia; Caucasus, but we have been unable to confirm ventral bristles on hind femur arising from this identification. slightly developed tubercles; hind Ta! straight In Zetterstedt's collection at Lund there were or with slight basal curvature; Ta4 setae straight three instances of a male and female placed on or with slight curve. the same pin. On the label of one of these, Wing: slightly infuscated with veins of cells Zetterstedt specifically noted that they were ri, r2+3 darkly pigmented. caught in copula. We were therefore able to Abdomen: dark brown. confidently associate the sexes. Genitalia (Fig. 10C): gonostylus with outer Little can be said about the bionomics of this margin evenly curved, tapering gradually species. Zetterstedt (1850) stated that males for basal half, with swollen, rounded apex; swarm near marshes in Lund, but unfortunately paramere (Fig. 10B) with apical half markedly his identifications of S. morio included both this swollen, apex rounded (but shrivelled in some); species and S.femorata. aedeagus (Fig. 10A) with lateral prongs, di- Goetghebuer (1936b) suggested that S. morio rected anterolaterally, distal portion a simple, was restricted to eutrophic lentic habitats in gradually tapering projection, about twice as southern Belgium but we were unable to confirm long to markedly longer than lateral prongs, this identification from his material. extreme apex directed ventrally. Labels on some specimens we examined noted Female adult. Descriptive statistics: see the following: 'from saline meadow', 'coastal Revision of Holarctic species of Serromyia Meigen 201

flat' (from Bar, Yugoslavia), 'at stream', 'at (fool) may refer to the unusual appearance of light'. Two female specimens deserve special this species as compared to the other species mention; they have the intriguing statement Fabricius placed in Culex. 'reared from nest of mole' (from near Beacons- field, Bucks., U.K.). Serromyia pacifica Remm sp.n. Strenzke (1950) and Thienemann (1950) re- corded larvae of S. morio from lake-side mosses Diagnosis. Male: only Palaearctic species in Germany but we have been unable to confirm with an AR = 1.17, the scutum pruinose, the these identifications. One pupa from Strenzke's fore femur entirely dark brown, with one row of collection labelled 'Strenzke No. 246 Serromyia ventral strong bristles and 1—2 posteriorly, and femorata (?) morio?' is actually a specimen of the parameres rounded apically. Female: only S.atra (see under that species and under generic Palaearctic species with palpus dark brown, discussion). Strenzke (1950: 349) also recorded nearly entire fore tibia darkly pigmented, a specimen as S.morio (?) from Ploner Sees, scutum pruinose, mid coxa with strong, lateral but we have not been able to examine this bristles, and with a single elongate hind claw,

specimen and confirm the identification. longer than Ta5. Taxonomic discussion. The type of S.morio Description. Male adult. Descriptive statistics: has been destroyed and only the label is left see Tables 2—7. (Zimsen, 1964). The type locality was given by Head: dark brown; antennal flagellomere 10 Fabricius (1775) as 'Anglia' [= England] and with plume arranged in more than one whorl; was collected on 13 May. We have accordingly palpus dark brown. designated a specimen from Oxford, U.K., Thorax: dark brown; scutum pruinose. collected at the same time of year, as neotype. Legs: coloration pattern as indicated in Fig. The type of S. nudipennis could not be located 3D; strong bristles of femora, tibiae distributed and is probably also destroyed. Kieffer's (1913) as follows: ventrally, posteriorly on fore femur, rather general description of the female of this posteriorly on fore tibia, anteriorly, ventrally, species does not allow for a definite application posteriorly on mid femur, posteriorly on mid of the name to any known European species. tibia, ventrally on hind femur, dorsally on hind However, most of the characters he does record tibia; ventral bristles on hind femur arising from correspond to the concept of S. morio as recog- slightly developed tubercles; hind Ta! with slight nized here (i.e pale wings, elongate hind claw). basal curvature; Ta4 setae straight or with slight Kieffer (1913) recorded this species from Bitche curve. (France). We examined two females from Wing: slightly infuscated with veins of cells

France but these were somewhat damaged. We rt, r2+3 darkly pigmented. therefore are designating a neotype (male) from Abdomen: dark brown. the same place and date as the neotype of Genitalia (Fig. 10F); gonostylus with outer S.morio (from England). margin every curved, tapering gradually for We examined a male of morio identified by basal half, with swollen, rounded apex; Zetterstedt as Ceratopogon flavicornis which paramere (Fig. 10E) with apical half somewhat was a misinterpretation of Staeger's description. swollen, apex rounded (somewhat shrivelled); Ceratopogon flavicornis is actually a member of aedeagus (Fig. 10D) with lateral prongs, di- the Bezzia (syntypes examined). Simi- rected dorsolaterally, distal portion a tapering, larly, some specimens identified by Meijere, simple projection, markedly longer than lateral Schiner, Edwards and Goetghebuer as prongs, extreme apex directed ventrally. S.femorata were in fact S.morio. Goetghebuer Female adult. Descriptive statistics: see also misidentified a specimen of S. morio as Tables 8—13. Similar to male except for usual S.atra. Remm had misidentified a female as sex differences and as follows: S. ledicola. Head: mandible serrate. The single paralectotype of S.spinosipes from Legs: coloration pattern as indicated in Fig. Budapest (BMNH) is a female adult of S.morio. 4E; strong bristles of femora, tibiae distributed Material examined. Ill males and 129 as follows: present or absent ventrally on mid females. femur, ventrally on hind femur; claw of hind leg Derivation of specific epithet. The name morio single, elongate, with basal tooth. 202 A. Borkent and B. Bissett

Wing: a few macrotrichia restricted to apical Serromyia rufitarsis: Kieffer 1906: 65. margin. Serromyia gelida Kieffer 1925b: 429. Neotype, Genitalia (as in Fig. 12D): sternite 9 truncate here designated, 8 adult on microscope slide, medially, with anteromedial margin developed, labelled 'Serromyia gelida Kieffer, Neotype, pointed; two spermathecae, additional sperma- Latvia, Sivera Lake, E. Remm, 17—6—67 thecal duct terminating in pigmented apex. CNC No. 20250, Serromyia dipetala R. det. Distribution and bionomics. S.pacifica is H. Remm, Serromyia rufitarsis Det. A. known from eastern Siberia (Fig. 18C). Adults Borkent' (CNCI). New synonym. examined were collected from 25 May to 27 Serromyia bispinosa Goetghebuer 1936a: 321. June. Lectotype, here designated, 8 adult on micro- The specimens from the type locality were scope slide, labelled 'La Panne [=De Panne, collected in mire. The material from Yakut Belgium], 8, 16-6-36, M. Goetghebuer', A.S.S.R. was taken from a taiga bog. 'Serromyia bispinosa', 'R.I.Sc N.B. 18.073 Taxonomic discussion. The description pro- Coll. et det., M. Goetghebuer', '8 Type vided here is based on the single male and two Lectotype Goetghebuer, Serromyia rufitarsus females housed in the CNCI and a manuscript Det. A. Borkent' (ISNB); paralectotype description provided by Mrs E. Remm. labelled as for lectotype except with '9' S.pacifica will also be described in a separate (ISNB). New synonym. paper, currently in press (Academy of Sciences, Serromyia dipetala Remm 1965: 182. Holotype, Tartu) and written by H. Remm before his 8 adult, not seen, from Estonian S.S.R., untimely death in 1986. Mrs E. Remm (pers. Valga District, shores of Lake Yakhiyavr, 11 comm.) indicated that the species description July 1952 (IZBE); paratypes: ESTONIA could be included in the present revision, to S.S.R.: 111c?, 699 (not seen) from Valga, ensure that this work is inclusive, and conse- Vyru, Pyarnu, Tartu and Kharyu districts, 13 quently whichever paper appears first will pro- June to 28 July, 1951-61; LATVIA S.S.R.: vide the date and citation for S.pacifica. 13<3\ 69, Daugavpils and Kraslava districts, Types. Holotype, 8 adult, from Sakhalin 21-24 June 1961 (IZBE). New synonym. Island, vicinity of Juzhno-Sakhalinsk, Ceratopogon morio: authors, not Fabricius. 16.vi. 1970, mire (IZBE); paratypes: 6cJ, Staeger 1839: 598 (in part). 79 labelled as for holotype (IZBE, CNCI); U.S.S.R.: 39, Kunashir island, Juzhno-Kurilsk Diagnosis. Male: only Palaearctic species in vicinity, 27.vi. 1970 (IZBE); 19, Primorye which the parameres bifurcate apically. Female: Territory, NP 'Kedrovaya Pad', l.vi.1970 only Palaearctic species with a shiny scutum, a (IZBE); 19 , from Amur Territory, Klimoutzy, completely darkly pigmented mid femur and a 25.v. 1957 (ZMAS); 18, 49 , Yakut A.S.S.R., single elongate hind claw. Yakutsk vicinity, 14.vi.1968 (IZBE). Description. Male adult. Descriptive statistics: Derivation of specific epithet. The name see Tables 2—7. pacifica refers to the proximity to the Pacific Head: dark brown; antennal flagellomere 10 Ocean, where this species was collected. with plume arranged in more than one whorl; palpus dark brown. Thorax: dark brown; scutum bare of Serromyia rufitarsis (Meigen) new status pruinosity. Ceratopogon rufitarsis Meigen 1818: 83. Lecto- Legs: coloration pattern as indicated in Fig. type, here designated, 8 adult on microscope 4B; strong bristles of femora, tibiae distributed slide, labelled 'Lectotype Ceratopogon as follows: present or absent posteriorly on fore rufitarsis 8 Meigen, ex coll. Meigen 293/ femura (one bristle), present or absent pos- 40 (160b), prep. J. Clastrier, 1985 baume teriorly on mid femur, ventrally on hind femur, phenole, Museum Paris' (MNHN); paralec- dorsally on hind tibia; ventral bristles on hind totype 8 adult labelled as for holotype but femur arising from slightly developed tubercles; with '160a' instead of '160b' and identified as hind Tat straight or with slight basal curvature; Serromyia femorata (MNHN). Ta4 setae straight or with slight curve. Prionomyia rufitarsis: Stephens 1829: 238. Wing: slightly infuscated with veins of cells Ceratolophus rufitarsis: Kertesz 1902: 158. rt, r2+3 darkly pigmented. Revision of Holarctic species of Serromyia Meigen 203

Abdomen: dark brown. description by Kieffer (1925b) of a male with Genitalia (Fig. 11C): gonostylus with outer brilliantly shiny thorax and a single spine on the margin evenly curved, tapering gradually for fore femur could apply to either S.atra or basal half, with rounded to somewhat pointed S. rufitarsis. On the basis of the general descrip- apex; paramere (Fig. 11B) with apical portion tion of leg pigmentation, we have placed the bifurcate, the ventral arm slender, the dorsal, species here. However, the type locality of posteriorly directed arm tapering sharply to S.gelida was given as northern Norway (just point; aedeagus (Fig. 11 A) with lateral prongs, north of Tromsoe [=Tromso]) which is directed laterally, dorsolateral^ or slightly markedly farther north than any locality of anterolaterally, distal portion a tapering projec- S.rufitarsis (Fig. 19C) or S.atra (Fig. 18B). The tion about twice as long to markedly longer only species with far northern distributions are than lateral prongs, extreme apex directed S.femorata (Fig. 19A) and possibly S.morio ventrally. (Fig. 19B) and S.ledicola (Fig. 18A). These Female adult. Descriptive statistics: see species, however, have either a pruinose scutum Tables 8—13. Similar to male except for usual or have many spines on the legs. It may be that sex differences and as follows: S. rufitarsis does occur in northern Norway but Head: mandible serrate. only future collecting will confirm this. How- Legs: coloration pattern as indicated in Fig. ever, the name S.gelida now must be considered 5F; strong bristles of femora, tibiae distributed a synonym of S.rufitarsis. as follows: present or absent ventrally on mid Remm (1965) noted that his S.dipetala dif- femur, ventrally on hind femur; claw of hind leg fered from S.bispinosa in lacking strong bristles single, elongate, with basal tooth. (his spines) on the fore and mid femora but we Wing: very few macrotrichia restricted to found this character to be variable, with a few apical margin. strong bristles present in some individuals (in- Genitalia (as in Fig. 12D): sternite 9 trunc- cluding some of those identified by Remm as ate medially to somewhat truncate but with S.dipetala). anteromedial margin developed, pointed; two Goetghebuer (1922a) noted that the syntype spermathecae, additional spermathecal duct series of Ceratopogon rufitarsis consisted of two terminating in pigmented apex. males. The male paralectotype of S.rufitarsis is Distribution and bionomics. S.rufitarsis is actually a specimen of S.femorata. known from England to Estonia south to Although Goetghebuer (1936a) based his de- Belgium and Hungary (Fig. 19C). Adults have scription on only one male and one female of been collected from 26 May to 23 July. S.bispinosa, it was uncertain which represented Remm (1965) recorded specimens from a few the holotype. Accordingly we have designated additional localities from Estonia and Latvia (as the male as lectotype. S.dipetala), which are also probably members Material examined. Thirteen males and ten of this species. females. The only available bionomic information is Derivation of specific epithet. The name from Remm (1965) who noted that this species rufitarsis (red tarsi) refers to the red-yellow tarsi (as S.dipetala) was most plentiful from 1 to 10 reported for this species by Meigen (1818) (see July and could be found not only in shrubs and generic taxonomic discussion for comments on thickets near water but also in damp meadows reports of red coloration by previous authors). some distance from any larger body of water. Taxonomic discussion. Although we were un- Serromyia subinermis Kieffer able to examine any type material of S.dipetala, Dr H. Remm kindly sent other material of this Serromyia subinermis Kieffer 1919: 73 (as species. Both the specimens and the detailed variety of S.spinosipes). Neotype, here description by Remm (1965) leave little doubt designated, 8 adult, labelled 'Serromyia that this name is a junior synonym of S. rufitarsis. subinermis Kieffer, 8, Neotype, Ocsa, The material sent by Remm also allowed correct Hungary, lapret, 1965.V. 19, leg. Mihalyi' association of males with females. (HNHM). The type of Serromyia gelida Kieffer could Serromyia femorata: authors, not Meigen. not be located and is presumed to be lost. The Zetterstedt 1838: 822 (in part), 1850: 3665 204 A. Borkent and B. Bissett

(in part). Edwards 1926: 410 (in part). 4F, 5E; strong bristles of femora, tibiae dis- Goetghebuer 1934: 61 (in part). tributed as follows: 0—4 ventrally on fore femur, Serromyia morio: Edwards 1928: 173, not 0-2 ventrally on mid femur, 0—2 apically on Fabricius. mid tibia, ventrally on hind femur; claw of hind leg single, elongate, with basal tooth. Diagnosis. Male: only Palaearctic species with Wing: a few to many macrotrichia restricted the parameres curved subapically and with the to apical margin. pointed apex directed laterally. Female: only Genitalia (as in Fig. 12D): sternite 9 trunc- Palaearctic species with the palpus dark brown, ate medially to somewhat truncate but with scutum pruinose, wing slightly infuscated with anteromedial margin developed, pointed; two veins of cells rl5 r2+3 darkly pigmented, lacking spermathecae, additional spermathecal duct strong lateral bristles on the hind coxa, and with terminating in pigmented apex. a single elongate hind claw. Distribution and bionomics. S.subinermis Description. Male adult. Descriptive statistics: is known from England to Estonia south to see Tables 2-7. Corsica, Yugoslavia and Hungary (Fig. 19D). Head: dark brown; antennal flagellomere 10 Adults have been collected from 10 April to 5 with plume arranged in more than one whorl; July. palpus dark brown. Taxonomic discussion. We have applied the Thorax: dark brown; scutum pruinose. name S. subinermis to this species by designating Legs: coloration pattern as indicated in Figs a neotype. The type material of this species is 4C, D; strong bristles of femora, tibiae dis- almost certainly lost. tributed as follows: anteriorly, posteriorly on Kieffer's (1919) brief description of this fore femur, present or absent ventrally on fore species noted that the fore and mid femora of femur, posteriorly on fore tibia, present or the female had one or two spines while the male absent dorsally, anteriorly, posteriorly on mid lacked spines altogether. Our description of the femur, present or absent dorsally, ventrally on female fits Kieffer's concept but that of the mid femur, posteriorly on mid tibia, present or male does not. It may well be that Kieffer absent anteriorly, dorsally, posteriorly on mid incorrectly associated the sexes as he did for tibiae, anteriorly, dorsally, ventrally on hind S.spinosipes (see under S.morio and S.atra). femur, dorsally on hind tibia, ventral bristles on We prefer to retain a given European name hind femur arising from slightly developed where possible, rather than propose a new one. tubercles; hind Tai straight or with slight basal We have examined specimens identified by curvature; Ta4 setae straight or with slight curve. Edwards and Goetghebuer as S.femorata, which Wing: slightly infuscated with veins of cells in fact are members of S.subinermis. Edwards r1? r2+3 darkly pigmented. (1928) recorded S. morio from Corsica but it is Abdomen: dark brown. actually a specimen of S.subinermis. Genitalia (Fig. 11F): gonostylus with outer The association of the sexes of S.subinermis margin evenly curved, tapering gradually to is somewhat tentative. None of the males and pointed apex; paramere (Fig. 11E) with sub- females we examined were collected at the apical portion somewhat expanded dorso- same place and time. Males and females col- ventrally (expansion best seen in lateral or lected in Hejobaba (8 km W Leninvaros), posterior view), with pointed apex directed Hungary, in May and males and females ident- laterally or anterolaterally; aedeagus (Fig- 11D) ified by H. Remm as S. spinosipes from Estonia with lateral prongs, directed posterolaterally to were considered correctly associated members laterally, distal portion a simple, somewhat of this species. broadened projection, about twice as long to In addition, we are not certain that all the markedly longer than lateral prongs, extreme females placed here are conspecific. Although apex directed ventrally. some appear to be correctly associated with the Female adult. Descriptive statistics: see males, some may belong to another, unnamed Tables 8-13. Similar to male except for usual species. Further collecting and study are sex differences and as follows: required to clarify this problem. Head: mandible serrate. Material examined. Twenty-five males and Legs: coloration pattern as indicated in Figs forty-one females. Revision of Holarctic species of Serromyia Meigen 205

Derivation of specific epithet. The name additional spermathecal duct terminating in subinermis (somewhat unarmed) probably pigmented apex. refers to the lack of spines on the fore and mid Distribution and bionomics. S.tecta is known femora which Kieffer (1919) attributed to males from only from the type locality in the Federal of this species. Republic of Germany (Fig. 19D). The date of collection on the labels is given as 6.5.1935 and Dr R. Contreras-Lichtenberg (NHMW) (pers. Serromyia tecta Borkent sp.n. comm.) has informed us that this means 6 May Diagnosis. Male: only Palaearctic species 1935. This clarification was added to the labels lacking spines on fore and mid femora and on the slide material. tibiae, each of which is entirely dark brown, Taxonomic discussion. Males and females of AR = 1.00—1.03, pruinose thorax and with S. tecta are very similar to those of Nearctic parameres rounded apically and swollen in S.borealis and they may be conspecific. How- apical half. Female: only Palaearctic species ever, males have different antennal ratios. with small, equal hind claws and a pruinose Larger series of specimens from a variety of thorax. localities are needed to determine whether this Description. Male adult. Descriptive statistics: is due to geographical variation or is indeed see Tables 2—7. indicative of genetic discontinuity. Head: dark brown; antennal flagellomere 10 Types. Holotype, 8 adult on microscope with plume arranged in more than one whorl; slide, labelled 'Holotype Serromyia tecta palpus dark brown. Borkent, 8, Traun [Austria], Czerny, 6.5.1935, Thorax: dark brown; scutum pruinose. May 6, 1935' (NHMW); allotype on microscope Legs: coloration pattern as indicated in Fig. slide labelled as for holotype (NHMW); para- 3D; strong bristles absent on femora, tibiae types: 38,29 labelled as for holotype (NHMW, except ventrally on hind femur, present or CNCI). absent dorsally on hind tibia; ventral bristles on Derivation of specific epithet. The name tecta hind femur arising from slightly developed (disguised) refers to the long hidden identity of tubercles; hind Tax straight or with slight basal this European species. curvature; Ta4 setae straight or with slight curve. Wing: slightly infuscated with veins of cells Fossil species r1? r2+3 darkly pigmented. Abdomen: dark brown. Several species of fossil Serromyia have been Genitalia (Fig. 12C): gonostylus with outer described, all from Holarctic localities, and margin evenly curved, tapering gradually these shed some light on the history of the for basal half, with rounded, somewhat genus. We were fortunate in being able swollen apex; paramere (Fig. 12B) with apical to examine all of these, except one male of half somewhat swollen, with rounded apex S.spinigera (Loew). Szadziewski (1988) has (shrivelled in some); aedeagus (Fig. 12A) with recently described the Baltic amber fauna lateral prongs, directed laterally to postero- of Ceratopogonidae and included six taxa of laterally, distal portion a simple, slender pro- Serromyia. These are: S.anomalicornis (Loew), jection, markedly longer than lateral prongs, S.spinigera (Loew), S.succinea Szadziewski, extreme apex directed ventrally. S.polonica Szadziewski and two unnamed Female adult. Descriptive statistics: see species (sp. A and B). His excellent and com- Tables 8—13. Similar to male except for usual prehensive study provides descriptions of these sex differences and as follows: species and we add here only a few further Head: mandible serrate. details. Because of the good preservation and Legs: coloration pattern as indicated in Fig. presence of some important character states, 5D; claws of hind leg equal, small. we have given names to Szadziewski's (1988) Wing: a few macrotrichia restricted to very species A and B. apical margin. Genitalia (as in Fig. 12D): sternite 9 some- Serromyia anomaiicornis (Loew) what truncate medially but with anteromedial margin developed, pointed; two spermathecae, Ceratopogon anomaiicornis Loew 1850: 30. 206 A. Borkent and B. Bissett

Types as indicated by Szadziewski (1988). Distribution and bionomics. S.colorata is Serromyia anomalicornis: Szadziewski 1988: known from compression fossils from the type 142. locality at Rott, Siebengebirge, Rheinland, Federal Republic of Germany. Although earlier Diagnosis and description. As indicated by thought to be Oligocene in age the Aquitanian Szadziewski (1988). Additional character states epoch is now dated as Miocene (Szadziewski, as follows: male antennal flagellomere 10 setae pers. comm.). in a single whorl; male, female with Ta4 with Taxonomic discussion. The above description straight setae. is based on the examination of the holotypes of Distribution. From Baltic amber. the species listed above and the single female Material examined. One male and one female. described by Statz (1944: 149) as Serromyia sp. Derivation of specific epithet. We were un- We were unable to find any significant differ- certain why the name anomalicornis (unusual ences between these specimens, based on the horn) proposed by Loew (1850) was applied to character states used in recognizing extant this species. species of Serromyia. We accordingly consider all to be conspecific. Statz (1941) provided the names Serromyia Serromyia colorata Statz spinofemorata and Serromyia austera under Serromyia colorata Statz 1944: 150. Holotype photographs of these, but failed to give any 9, both halves of split rock present, mounted diagnostic features. The names are therefore on wooden block, labelled 'Serromyia nomina nuda. However, the descriptions by colorata Stz. Original!', '3527', 'Serromyia Statz (1944) do validate these names. Sphon colorata Sz. nov. sp.' (LACM). (1973) incorrectly considered the earlier naming Serromyia austera Statz 1944: 150. Holotype 9, by Statz (1941) to invalidate his later use of the both halves of split rock present, mounted on names (see ICZN Art. 23:m). wooden block, labelled 'Serromyia austera Statz (1944) stated that the antennae of Stz. Original!', '3526', 'Serromyia austera S. colorata, S.austera and S.spino femorata each Sz.', 'Serromyia colorata Det. A. Borkent' had 14 segments but each actually has 13 (LACM). New synonym. flagellomeres. The macrotrichia visible at the Serromyia spinofemorata Statz 1944: 151. wing apex reported for S. colorata were actually Holotype 9 , only one half of split rock pres- those at the very margin. ent, mounted on wooden block, labelled We consider some of the slight colour differ- 'Serromyia spinofemorata Statz Original!', ences noted by Statz (1944) to be artefacts of '3528', 'Serromyia spinofemorata Sz.', preservation. 'Serromyia colorata Det. A. Borkent' Material examined. Four females. (LACM). New synonym. Derivation of specific epithet. The name colorata (colour) presumably refers to the dif- Diagnosis. Female: apical portion of fore ferences in color Statz (1944) attributed to this femur and tibia pigmented, hind claw elongate species. with HC/Ta5 = 1.09-1.38. Description. Male adult. Not known. Serromyia polonica Szadziewski Female adult. Legs: coloration pattern with apical portion of fore and mid femora, basal Serromyia polonica Szadziewski 1988: 135. portion of fore and mid tibiae darkly pigmented Types as indicated by Szadziewski (1988). (Fig. 13D); strong bristles absent on femora, tibiae except present ventrally on hind femur Diagnosis and description. As indicated by (Figs 13A, C); hind femur length = 0.86-1.07 Szadziewski (1988); additional character state mm; claw of hind led single, elongate (Figs 13B, as follows: Ta4 with straight setae. F), with basal tooth, HC/Ta5 - 1.09-1.38. Distribution. From Baltic amber (Eocene). Wing: wing venation not discernible. Taxonomic discussion. We could not confirm Genitalia (as in Fig. 13D): sternite 9 truncate the presence of the prescutal pits as reported by medially, two spermathecae present (additional Szadziewski (1988). If well developed, they spermathecal duct not discernible). would be unique in the genus. Revision of Holarctic species of Serromyia Meigen 207

Material examined. Two females (all the type Type. Holotype, 8 adult in amber, in paper material). envelope labelled 'Diptera Cera- Derivation of specific epithet. The name topogonidae Serromyia sp. A, Szadziewski 1988 polonica presumably refers to the type locality 8. MZ 14972, Serromyia sinuosa Borkent' in Poland. (MZW). Derivation of specific epithet. The name sinuosa (windings) refers to the sinuate setae Serromyia ryszardi Borkent sp.n. found on the fourth tarsomeres of this species. Serromyia sp. B Szadziewski 1988: 139. Serromyia spinigera (Loew) Diagnosis and description. Described by Szadziewski (1988: 139) as Serromyia sp. B. Ceratopogon spiniger Loew 1850: 30. Types as Additional character states as follows: antennal indicated by Szadziewski (1988). flagellomere 10 with plume arranged in a single Serromyia spinigera: Kieffer 1906: 2. whorl; strong bristles of femora, tibiae distri- Szadziewski 1988: 140. buted as follows: anteriorly, ventrally, posteriorly Ceratopogon elongatus Meunier 1904: 242. on fore femur, anteriorly, dorsally, posteriorly Types as indicated by Szadziewski (1988). on fore tibia, anteriorly, ventrally, posteriorly Serromyia elongatus: Szadziewski 1988: 140. on mid femur, dorsally, posteriorly on mid tibia, anteriorly, dorsally, ventrally on hind Diagnosis and description. As indicated by femur, 2—3 rows dorsally on hind tibia; aed- Szadziewski (1988). Additional character state eagus with trifid apex; Ta4 with straight setae. as follows: Ta4 with straight setae. Distribution. From Baltic amber (Eocene). Distribution. From Baltic amber (Eocene). Taxonomic discussion. The trifid aeadeagus Material examined. Two females. of S.ryszardi is somewhat similar to that of Derivation of specific epithet. The name several extant species in the Holarctic (see Figs spinigera (spine bearer) presumably refers to 8A, D, 9D), with the lateral prongs and the the noted spines on the hind femur of this apex directed in a posterior direction. species. Type. Holotype, 8 adult in amber, in paper envelope labelled 'Diptera Nematocera Cera- Serromyia succinea Szadziewski topogonidae Serromyia sp. B, Szadziewski 8. MZ 16110, Serromyia ryszardi Borkent' (MZW). Serromyia succinea Szadziewski 1988: 136. Derivation of specific epithet. The name Types as indicated by Szadziewski (1988). ryszardi is named in honour of the many contri- butions that Ryszard Szadziewski had made to Diagnosis and description. As indicated by our understanding of ceratopogonid systematics Szadziewski (1988). and, in particular, his comprehensive work on Distribution. From Baltic amber (Eocene). Baltic amber material. Taxonomic discussion. Flagellomere 10 of the male could not be seen clearly. Both male and females had only straight setae on ta . Serromyia sinuosa Borkent sp.n. 4 The male was correctly reported to have Serromyia sp. A Szadziewski 1988: 138. a few macrotrichia at the wing tip (discussed below under 'Phylogeny'). Diagnosis and description. Described by Material examined. One male and two females Szadziewski (1988: 138) as Serromyia sp. A. (all type material). Additional character states as follows: antennal Derivation of specific epithet. The name flagellomere 10 with plume arranged in a single succinea (amber) clearly refers to the state of whorl; Ta4 with sinuate setae. preservation of this species. Distribution. From Baltic amber (Eocene). Taxonomic discussion. The sinuate setae on Discussion of fossil material the fourth tarsomere are unique amongst the Baltic amber and extant Holarctic species of Our examination of many specimens of extant Serromyia. species as well as of available fossils of extinct 208 A. Borkent and B. Bissett species allows for some improvements in the name available until such time as further key given by Szadziewski (1988). In particular, specimens become available for study and a our studies show that the ratio of the second neotype can be designated. radial cell to the first is highly variable and a poor tool for recognizing extant species. This suggests that the same would be true for the Phylogeny fossil species. The suggested changes replace A fundamental question to the interpretation the use of that character in the key and are as of phylogenetic relationships within Serromyia, follows: concerns the monophyly of the genus itself. Throughout the course of this study the presence of a swollen hind femur bearing a Couplet 6 ventral patch of strong bristles arranged as a Fourth tarsomerc with both straight and one pair of single row of 1—3 bristles basally, forming 2 sinuate setae S.sinuosa rows at mid length and scattering distally into Fourth tarsomerc with only straight setae 7 an indistinct pattern up to 4 bristles wide, was interpreted as a derived character state. This character state appears to be unique within the Couplet 8 Ceratopogonidae and . Fore and mid femur with strong bristles anteriorly, Wirth & Grogan (1988), however, recently ventrally and posteriorly S.ryszardi described the new monotypic genus Metacan- Fore and mid femur with only a few strong bristles thohelea which exhibits this same character state. ventrally S.succinea On the basis of this modification, we suggest that these two together form a monophyletic Because S. colorata is known as a compression group (Fig. 20). fossil, it was not included in the key by Wirth & Grogan (1988) suggested several Szadziewski (1988). If keyed out, it would run characters which may serve to distinguish to S.succinea. Although S.colorata has darker Metacanthohela from Serromyia. Our study leg pigmentation restricted to the apex of the indicates that many of these also occur in at fore femur and tibia and S.succinea females least some species of Serromyia. The proposed have completely dark fore legs, it is unlikely distinguishing character of Metacanthohelea, that this can be used to distinguish them followed by the character states within Virtually all Baltic amber ceratopogonids lack Serromyia are given below: contrasting pigmentation and this may be an artefact of preservation (Szadziewski, 1988). — slightly arcuate hind tibia: indistinguishable from We can suggest no other distinguishing feature. that of S.reyei, S.maculipennis, S.mangrovi, S.silvatica, Another fossil, described by Heyden (1870) S.stuckenbergi, S.nocticolor. The degree of bend of as Ceratopogon alpheus, is probably a member the base of the hind tibia seems strongly correlated with the thickness of the hind femur. Thicker hind of Serromyia. The male specimen was collected femora are associated with a stronger bend at the from the brown coal deposits at Rott, base of the tibia. Siebengebirge, Federal Republic of Germany. The figure clearly shows plumose antennae and — reduced number of hind femur spines: within markedly swollen hind femora and we know of Serromyia, the number of spines (here called strong no other Nematocera with such a combination bristles) is generally correlated with the size of the hind femur. However, the extent to which the hind of characters. Some and Schizohelea femur bears strong bristles does vary. Female S.reyei, also have swollen hind femora but not to such a S.silvatica and S.mangrovi have the same or a smaller degree. We therefore recognize this species as a percentage of the hind femur covered with strong new combination: Serromyia alpheus (Heyden). bristles when compared with M.cogani. Unfortunately, the type was reported as lost The male of M.cogani has the distal 0.39-0.43 of (Szadziewski, 1988), and we too have tried the hind femur bearing spines. Within Serromyia the without success to locate it at the BMNH and least amount of the hind femur bearing strong bristles OXUM. It is possible that this male was is exhibited by S.reyei with a minimum of 0.58. conspecific with the females described as Otherwise, all Serromyia have an even more exten- S.colorata. However, we prefer to leave the sively spinose femur. Revision of Holarctic species of Serromyia Meigen 209

— female with equal hind claws: also present in The first suggests that the plesiomorphic state S.silvatica, S.atra, S. crassifemorata, S. borealis, is the total lack of strong bristles (as in most S.tecta. However, in each of these Serromyia species, Ceratopogonidae), developing to a state where each claw does not bear an inner tooth, which is strong bristles are restricted to the distal portion present in Metacanthohelea coqani. of the hind femur (as in Metacanthohelea), and — female with relatively short hind fourth ultimately to the extensive strong bristles of tarsomere: we failed to find any significant differences Serromyia. This hypothesis would suggest between the hind fourth tarsomeres of Metacan- that Metacanthohelea is the sister group of thohelea and species of Serromyia. Serromyia. — eyes broadly separated: also present in S.reyei, The second hypothesis suggests that the S.mangrovi, S.stuckenbergi and S.nocticolor. plesiomorphic condition is a single, but exten- — male antenna with flagellomeres 1-8 fused: sive row of strong bristles (as in Echinohelea flagellomeres fused in S.mangrovi and S.nocticolor or Macfie, some Austrohelea Wirth & Grogan, at least partially fused basal flagellomeres in all some Fanthamia de Meillon and some Holarctic species other than S.nudicolis. ), which develop into the more complex pattern typical of most species of — parameres fused: also fused in S.reyei, S.festiva and S.esaki. Serromyia. This suggests that Metacanthohelea cogani is merely an autapomorphic member of — sensory pit on palpal segment 3: also present in Serromyia, with a somewhat reduced distri- male and female S. agathae and 5. nocticolor and in at bution of strong bristles on the hind femur. least the male of S.festiva. In our opinion, the first hypothesis seems — radically different genitalia: although the male most likely and we therefore recognize Meta- genitalia of Metacanthohelea are somewhat different canthohelea as a valid genus (Fig. 20). We are from those of most Serromyia species, they are similar unable to suggest an apomorphy for Meta- to those of S.reyei. In addition, some Serromyia also canthohelea, but, considering the genus is have rather different genitalia when compared monotypic, it must be monophyletic. to those in the rest of the genus (e.g. S.esakii, S. mangrovi). In spite of a diligent search for character states which might be useful in interpreting the From the above comparisons, it appears that phylogenetic relationships between species of the only possible distinguishing character of Serromyia, only a few clues were discovered. Metacanthohelea is the reduced extent to which In all extant Holarctic species of Serromyia, the male hind femur bears strong bristles. In other than S.mangrovi, the tenth flagellomere addition, re-examination of the original material exhibits a series of whorls of elongate bristles. (Wirth, pers. comm., and ourselves) indicates In all other Ceratopogonidae, including all that the female claws are distinctive. They are Serromyia examined from the Oriental, bent sharply at their base and with the distal Australian and Afrotropical Regions, the tenth portion straight (Fig. 16C), in comparison to flagellomere has a single whorl of elongate species of Serromyia, where the base of the bristles, similar to that of the preceding fla- claws are either straight or evenly curved and gellomeres. The multi-whorled condition is the distal portion of the claw is always curved therefore logically interpreted as a synapo- (Figs 16D—G). In addition, the claws of the morphy (Fig. 20). This character state is also hind leg of M.cogani, although equal and shorter reflected in the ratio of antennal flagellomere than the fifth tarsomere, bear a well-developed 10/11 (Table 3), where S.mangrovi has a single inner tooth. In those Serromyia species which whorl and a low ratio and all other species have have equal hind claws, the inner tooth is lacking. a higher ratio and have more than one whorl. Of the above character states we can inter- Considering the monophyly of the Holarctic pret only the degree to which the male hind species (except S.mangrovi), another character femur bear strong bristles for phylogenetic an- state may give further resolution. All Palae- alysis. Outgroup comparisons indicate that most arctic species (except S.mangrovi) and several genera of Ceratopogonidae lack thick strong Nearctic species exhibit lateral prongs on the bristles on the hind femur but that some bear a aeadeagus. This tripartite condition is also single row of thick spines. Two alternate hy- present in various configurations, in some potheses are therefore possible. African species: S.agathae, S.neethlingi, 210 A. Borkent and B. Bissett

•2 2. £ o § t ^ 3s Qj -S? Co .y o o s o V-o" co: <35 OJ CO •S CD c o s o I 2 I 0j c o CO c + CO Co Co

aedeagus lacking side prongs

flagellomere 10 with several whorls of long setae

male with >0.58 of hind femur with strong bristles

2-4 rows of strong bristles on hind femur

Fig. 20. Cladogram showing relationships within Serromyia and its sister lineage. Black rectangles and accompanying descriptions represent synapomorphic character states.

S.meiswinkeli, S.nocticolor and S.stuckenbergi. presence of long claws in the genus and similar This includes all those Serromyia south of the genera such as Monohelea and Stilobezzia. Holarctic in which the body coloration is entirely Although the presence of short claws is probably dark brown and for which the male is known. apomorphic within Serromyia, incompatibility This may allow for interpretation of the two with other character states suggests that this character states found within the Holarctic character state is susceptible to homoplasy and lineage where the aedeagus either has lateral argues against such a grouping. S.silvatica, an prongs or is lacking these. Using the African African species known only from the female, species as the outgroup, those species lacking has short hind claws. It is unlikely that the male lateral prongs in the Nearctic would form a will have the synapomorphy of possessing more monophyletic group: S. barberi, S. crassi- than one whorl of setae on flagellomere 10, a femorata and S. nudicolis (Fig. 20). synapomorphy grouping all Holarctic species Although based on phenetic similarity, it (except S.mangrovi), including four species seems likely to us that S. morio and S.atra are which have short hind claws: S.atra, S.borealis, sister species. The male genitalia of these two S.tecta and S.crassifemorata. Considering the species were inseparable. overall morphological similarity between Szadziewski (1988: 142) has suggested that S.borealis and S.tecta, these two are probably those species where the females have short, sister species. However, as argued above, equal hind claws may form a monophyletic S. crassifemorata is more closely related to two group, which he called the crassifemorata group. long-clawed species than to other short-clawed This hypothesis seems unlikely to us. It does species. In addition, also as suggested above, seem clear that the character state of having the short-clawed S.atra is probably most closely short claws is derived from a plesiomorphic related to S.morio, a species with an elongate longer, single claw, based on the widespread hind claw. Finally, two of the fossil species Revision of Holarctic species of Serromyia Meigen 211

S.spinigera and S.anomaiicornis have short hind Serromyia males. However, this character state claws and assuming these are correctly associ- cannot be interpreted as synapomorphic, as ated with the males, definitely do not belong to male wing macrotrichia also occur in a number the clade including the extant Holarctic species of other ceratopogonid genera. (three species of which have short claws). We Previous workers have used some characters conclude, therefore, that short hind claws have states to recognize new taxa (particularly genera) evolved independently at least three times which show homoplasy within Serromyia. If amongst the extant Holarctic lineage and that it Serromyia is indeed monophyletic, this may is a poor indicator of relationship. The question indicate that those character states are sus- may be further tested by discovery of the male ceptible to homoplasy in other groups of cera- of S.silvatica and further resolution of phylo- topogonids as well and that they should be genetic relationships amongst the Holarctic interpreted with caution. The character states lineage. are as follows: separate or fused male flagello- As noted above, S.mangrovi is not part of meres; terminal three or four male flagello- clade formed by all other Holarctic species. meres elongate; presence or absence of sensory This, its distinctive morphology (as compared pit on third palpal segment; female mandible to Holarctic species), and its presence in the vestigal or well developed; distance between Sinai, may indicate an Afrotropical or Oriental eyes; presence or absence of pruinosity on scutum; phylogenetic connection for this species. base of M2 indistinct or well developed; presence Our examination of fossil material provided or absence of macrotrichiae on male or female some clues about the history of the genus. All wings; presence or absence of strong bristles on males of the fossil species had a single whorl of legs; presence or absence of sinuate setae on setae on flagellomere 10, indicating that fourth tarsomeres; short and equal or elongate they were not members of that clade which and single hind female claws; straight or basally includes all extant Holarctic species (except curved hind first tarsomere; presence or absence S. mangrovi). Either the Holarctic species of stout spine near base of hind first tarsomere; diversified since the Eocene (Baltic amber age) female with sternite eight bilobed or completely or they evolved elsewhere. cleft; separate or fused parameres; 1—3 func- S. ryszardi was unique in the fossil material in tional spermathecae. exhibiting 2—3 rows of strong dorsal bristles on Debenham (1987) recently described the new, the hind tibia. The only extant Serromyia species monotypic genus Chimaerohela and suggested which has a similar condition is S.nocticolor that it was closely related to Serromyia. This from South Africa. This character state is prob- conclusion was based on shared similarities ably a valid synapomorphy. Only some members between Chimaerohelea and some species of of Echinohelea have a similar condition. Serromyia. These stated similarities require Two other clues indicate that the Baltic am- comment, based on our study of Serromyia. ber Serromyia may not be immediately related The following gives the character state which to the assemblage of extant Holarctic species. Debenham (1987) suggested was shared by The sinuate setae on the fourth tarsomere of Chimaerohelea and various Serromyia, followed S.sinuosa (Baltic amber) is shared only with by our observations: some more southerly members of the genus: S.reyei (Australasian), S.stuckenbergi (Afro- — fusion of the veins between cells ^ and r2+3 present in S.barberi, male S.reyei, S.femorata and tropical) and S.agathae (Afrotropical). It is possibly S.nocticolor: our examination showed that uncertain, however, which character state is no Serromyia species showed such fusion cxcept as derived. Both conditions of sinuate and straight intraspecific variation in a few species (S.nudicolis, setae on the fourth tarsomere are widespread S.femorata, S.atra). throughout the . — a single spermatheca present in S.aethiopiae: this A second character state is shared by is the only species of Serromyia with a single S.succinea (Baltic amber) and S.agathae (Afro- spermatheca. However, the character state appears tropical). Males of both have a few macrotrichia to be homoplastic in numbers of other genera of at the wing apex. Although the presence of Ceratopogonini ( Kieffer (1-2), wing macrotrichia are widespread on the wings Kieffer (1-2), Ceratoculicoides Wirth of females, they are not present on any other & Ratanaworabhan (1-2), Kolenohelea de Meillon 212 A. Borkent and B. Bissett

& Wirth (1—2), Macrurohelea Ingram & Macfie (1 — distributions to be confidently interpreted. The 2), Stilobezzia Kieffer (1-3)) and is probably a poor only Holarctic species, S.ledicola, is widespread indicator of relationship. in the Nearctic but appears to be absent from

— wing fold extending basally from point where M2 most of the area west of the continental divide curves toward M^ we confirm that S.aethiopiae is the in more southerly latitudes. only Serromyia with a barely discernable, slightly Within the Palaearctic, all the European pigmented line extending basally. species which were well represented in collec- — male flagellomeres 4-11 fused in S.dipetala tions, seem to be broadly distributed in central (named S.rufitarsis here), possibly S.nocticolor, and . In southern Europe, however, species some other genera of Ceratopogonini: we found fused appear to be restricted to mountainous regions. flagellomeres present in S.mangrovi, S.nocticolor and Although S.femorata is the only species defi- at least partial fusion in all Holarctic species other nitely known from northern Fennoscandia, it than S.nudicolis. seems likely that at least S.morio (based on its — aedeagus similar to S.femorata, S.pendleburyi, presence in Scotland) and S.ledicola (based on S.dipetala (named S.rufitarsis here): we failed to see its northern distribution in North America) will distinctive similarities between Chimaerohelea and also be found there. any species of Serromyia. Szadziewski (1988) has suggested a European origin for the genus Serromyia. We can see no Of the characters discussed by Debenham logical basis for such a conclusion. We recognize (1987), the only character state which may that zoogeographic interpretation must be based possibly be interpreted as a synapomorphy is on an understanding of the phylogenetic re- the dark line extending basally from M2. We lationships within the group under consider- could find no other Ceratopogonini with such a ation. Until further interpretation of the character state. However, this would indicate relationships between the known species is that the monopoly of Serromyia plus Metacan- available, it will be impossible to identify the thohelea would be in doubt. We consider this to early lineages of Serromyia and thereby suggest be unlikely and that the above similarities are where the genus may have originated. all due to homoplasy. Szadziewski (1988) also interpreted his We are unable, however, to present an 'crassifemorata group' as exhibiting a recent alternative hypothesis of the phylogenetic Euro-North American distribution. As indi- position of Chimaerohelea. Further research is cated above, however, evidence suggests that require to interpret the position of this and this group is polyphyletic (grouped on the basis many other genera of Ceratopogonini. of parallelism). As such, no zoogeographic interpretation could be validated. The fossil data indicate that the extant Zoogeography Holarctic lineage was probably not present in The above phylogenetic interpretation is so Europe during the Eocene. We are inclined to sketchy that it is presently impossible to inter- think that the extant Holarctic species, as a pret the historical zoogeography of the extant monophyletic group, are most closely related to Holarctic species. However, some distributions those species from Africa which are darkly allow for some general zoogeographic state- coloured. However, there is no logical evidence ments to be made. to support this at present, other than some of The distributions of all species of Serromyia the unpolarized shared similarities between in the Nearctic are restricted to temperate Holarctic species (other than S.mangrovi) and habitats. The southern extensions are otherwise some of the African species noted above. present only in the mountainous regions. No Serromyia are known from the Neotrophical Acknowledgments Region. Within the Nearctic, S.nudicolis and S.crassi- This study was the result of 4 years of investi- femorata are restricted to eastern North gation. A few years ago, when the senior author America. S. barberi is present only west of was new to the study of Ceratopogonidae, Dr the continental divide while the remaining Willis W. Wirth gave generously of his time and species are too poorly represented for their expertise to help in initiating my studies. He Revision of Holarctic species of Serromyia Meigen 213

already had a draft key of the Nearctic species thorough reviews of this manuscript: Dr William of Serromyia (including S.nudicolis) as well as L. Grogan, Dr Evert E. Lindquist and Dr some notes on the species. He unstintingly gave Michael Sharkey, Dr Ryzard Szadziewski, Dr these to us. We express our thanks to him for Richard J. Vockeroth and Dr Willis W. Wirth. his magnanimous approach and support. Their suggestions and criticisms greatly im- Leo Forster mounted on microscope slides proved the text. nearly all the specimens used in this study. His Finally, the senior author thanks his wife remarkably clear and carefully prepared mounts Annette and his three children for sharing sev- added immeasurably to the information gath- eral field trips and assisting in many of the tasks ered during this work. He also helped with demanded by these excursions. His son Chris some field collecting. Without his assistance this also helped with an attempt to interpret strong study would not have compiled the information bristle variation in S.barberi males. it has. His retirement at the end of 1988 was sorely felt! We also appreciate the assistance of the staff References of the Electron Microscope Unit of the Cell Arnett, R.H. & Samuelson, G.A. (1986) The Biology Research Centre, Agriculture Canada. and Spider Collections of the World, 220pp. E. J. Barry Flahey drew and inked all the genitalia Brill/Flora and Fauna Pub., Gainesville, Florida. drawings. His skills in illustrating these small Becker, P. (1961) Observations on the life cycle and creatures are much appreciated. immature stages of Culicoides circumscriptus Kieff. Ms Mary Dillon kindly identified some (Diptera, Ceratopogonidae). Proceedings of the chironomids which were recorded as prey. Royal Society of Edinburgh (B), 67, 363-386, We also extend our thanks to Mrs Evi Remm, Plates 1,2. who kindly provided advice on how to deal with Boorman, J. & Rowland, C. (1988) A key to the S.pacifica Remm. genera of British Ceratopogonidae (Diptera). Our thanks also extends to all those curators Entomologist's Gazette, 39, 65-73. Borkent, A. (1984) The systematics and phylogeny of who lent specimens of Serromyia: Dr R. the Stenochironomus complex (Xestochironomus, Abraham (ZMUH), Dr P. H. Arnaud (CASC), Harrisius, and Stenochironomus) (Diptera: Mr B. Brugge (ZMAN), Dr M. Chvala Chironomidae). Memoirs of the Entomological (KUPC), Dr J. Clastrier (MNHN), Dr R. Society of Canada, 128, 269pp. Contreras-Lichtenberg (NHMW), Dr P. S. Clastrier, J. (1960) Notes sur les Ceratopogonides. X. Cranston and Mr B. Townsend (BMNH), Mr Ceratopogonides de la Republique du Congo (2). R. Danielson (MZLU), Dr J. C. Delecolle Archives de I'lnstitut Pasteur d'Algerie, 38, (MZSF), Dr A. Dely-Draskovits (HNHM), Dr 258-298. N. L. Evenhuis (BPBM), Dr W. L. Grogan Clastrier, J. (1963) Notes sur les Ceratopogonides. (Maryland), Dr P. Grootaert (ISNB), Dr P. XVIII. Especes du genre Stilobezzia Kieffer ou Apparentees de la Region Palearctique. Archives Havelka (PEHC), Dr J. K. Liebherr (CUIC), de I'lnstitut Pasteur d'Algerie, 41, 41-68. Dr B. Lindeberg (UZMH), Dr L. Lyneborg Clastrier, J. & Wirth, W.W. (1961) Notes sur les (ZMUG), Dr G. C. McGavin (OXAM), Ms. Ceratopogonides. XIII. Ceratopogonides de la K. C. McGiffen (INHS), Ms P. 1. Persson region ethiopienne. Archives de I'lnstitut Pasteur (NHRS), Dr W. W. Wirth and Dr B. V. d'Algerie, 39, 190-240. Peterson (USNM), Dr F. Reiss (ZSMC), Dr H. Coquillett, D.W. (1900) Papers from the Harriman Remm (IZBE), Mr L. Saul (LACM), Dr R. T. Alaska Expedition. IX. Entomological results (3): Schuh (AMNH), Dr R. O. Schuster (UCDC), Diptera. Proceedings of the Washington Academy Dr R. Szadziewski (Gdansk). In addition Dr of Sciences, 2, 389-464. G. C. McGavin gave us substantial help with Debenham, M.L. (1970) Australasian Cerato- interpreting otherwise impossible British place pogonidae (Diptera, Nematocera). Part XIV: The genus Serromyia Meigen. Proceedings of the names, Ms Evi Remm provided much appreci- Linnean Society of New South Wales, 94, 160-165. ated help with Latvian, Estonian and Lithuanian Debenham, M.L. (1987) Chimaerohelea, a new genus localities, Dr L. Papp interpreted Hungarian of Ceratopogonidae (Diptera) from North place names and Dr P. Grootaert kindly did the Queensland. Invertebrate , 1, 801-806. same with difficult Belgian localities. Delecolle, J.-C. & Braverman, Y. (1987) Description We thank the following who provided de Serromyia mangrovi n.sp. du Sinai (Dipt. 214 A. Borkent and B. Bissett

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Band, 18, 1-202. Agricultural Research Service. Thomson, C.G. (1869) Diptera species novas de- Wirth, W.W. (1973) Family Ceratopogonidae. Pp. scripsit. Pp. 443 - 614, PI. 9, in Kongliga Svenska 346-388 in A Catalog of the Oriental Region, fregatten Eugenies resa omkring jorden. Pt 2: Volume 1. Suborder Nematocera, 618pp. Univer- Zoologi. 1. Insekter, 617pp., 9 pis. Stockholm, sity Press of Hawaii, Honolulu. '1868'. Wirth, W.W. & Grogan, W.L. (1981) Natural History Thomsen, L.C. (1937) Aquatic Diptera Part V. of Plummers Island, Maryland. XXV. Biting midges Ceratopogonidae. Memoirs of the Cornell Uni- (Diptera: Ceratopogonidae). 3. The species of the versity Agricultural Experiment Station, 210,57—80, tribe Stilobezziini. Bulletin of the Biological Society Pis. 10-18. of Washington, 5, 1 — 102. Tokunaga, M. (1940) Ceratopogonidae and Chiro- Wirth, W.W. & Grogan, W.L. (1988) The predaceous nomidae from the Micronesian Islands. Philippine midges of the World (Diptera: Ceratopogonidae; Journal of Science, 71, 205—226. Tribe Ceratopogonini). Flora and Fauna Handbook Westwood, J.O. (1840) Order XIII. Diptera Aristotle No. 4, xv + 160pp. E.J. Brill, Leiden. (Antliata Fabricius. Halteriptera Clairv.). Yu, Y-X. (1978) Chapter III. The identification Pp. 125-128 (= signature I, part), 129-144 of important biting midges of China [in Chinese] (= signature K), 145-158 (= signature L) in An pp. 178-224, Pis. 16-22. Manual of Important Introduction to the modern classification of insects. Chinese Insect Vectors of Disease. People's Synopsis of the genera of British insects, 158pp. Health Publishing House, China. London. Zetterstedt, J.W. (1838) Diptera Scandinaviae. Sect. Williston, S.W. (1907) Dipterological Notes. Journal 3: Diptera. Pp. 477-868 in Insecta Lapponica vi + of the New York Entomological Society, 15, 1—2. 1140pp. Lipsiae [=Leipzig]. Winnertz, J. (1852) Beitrag zur Kenntniss der Gattung Zetterstedt, J.W. (1850) Diptera Scandinaviae. Ceratopogon Meigen. Linnea Entomologica, 6, Disposita et descripta, Vol. 9, pp. 3367-3710. 1-80. Lundae [=Lund], Wirth, W.W. (1952) The Heleidae of California. Zihali-Sebess, G. (1940) Magyarorszag Heleidai. University of California Publications in Ento- (Heleiden aus Ungarn). Folia Entomologia mology, 9, 95-266. Hungarica, 5, 10-133. Wirth, W.W. (1965) Family Ceratopogonidae. pp. Zimsen, E. (1964) The type material of I. C. Fabricius. 121-142 in Stone, A., Sabrosky, C.W., Wirth, Munksgaard, . W.W., Foote, R.H. & Coulson, J.R. Eds. A Catalog of the Diptera of America North of Mexico, 276 pp. United States Department of Agriculture, Accepted 27 October 1989 Systematic Entomology (1990) 15, 218

Erratum Vossbrinck, C.R. & Friedman, S. (1989) A 28s ribo- somal RNA phylogeny of certain cyclorrhaphous Diptera based upon a hypervariable region. Sys- tematic Entomology, 14, 417—431.

The tsetse species listed as Glossina simulans should be Glossina morsitans. 160 A. Borkent and B. Bissett

Teskey (1981) and detailed characters (i.e. some groups have bristles that are more chaetotaxy) as well as pupal characters follow markedly developed than in any species of Saunders (1924) and Lawson (1951). Serromyia. The strong bristles as used here to We use some terms in the keys and descrip- describe character states refer to those bristles tions that require further comment. Some adult which stand out from the remaining setae as Serromyia lack pruinosity on the scutum and in thicker and often appear to be more darkly such specimens the scutum is brilliantly shiny pigmented. Figs 1C, D indicate examples of and highly reflective in pinned material (the typical strong bristles on the fore and hind leg, humeral pits are not distinct from the surround- respectively. The lateral strong bristles on the ing shiny cuticle) and utterly lacking fine spicules hind coxa are shown in Fig. IB. Sometimes when viewed laterally in slide-mounted speci- strong bristles are broken off at the base and mens. Specimens which are reported to exhibit then the enlarged socket needs to be searched pruinosity on the scutum are somewhat dull for, for accurate interpretation of keys and in reflected light (so that the humeral pits ap- descriptions. pear as discrete, shiny patches). When slide Leg coloration is illustrated somewhat dia- mounted, such specimens have a short coat of grammatically. Leg pigmentation intensity dif- fine spicules visible in lateral view (Fig. 1A). fered between specimens and varied according We looked for pruinosity amongst the dorso- to preparation technique. Illustrations therefore central setae on the laterally mounted thorax are meant to show extent of pigmentation for a using interference contrast optics. given species with the intensity of pigmentation When one examines the brilliantly shining not necessarily comparable between species. scutum of a pinned specimen with a dissecting Parameres are described as either rounded or microscope, one can be confident of the lack of tapered apically. Rounded parameres are some- pruinosity; but when the scutum is dull, care times shrivelled and may appear pointed. How- must be taken. A somewhat dirty specimen may ever, in most such instances the two parameres appear dull when in fact it lacks pruinosity. In look different from one another. Parameres most of these instances the humeral pits would which are tapered apically are always of clearly also be dull. We generally preferred to examine defined and characteristic form, as illustrated in slide-mounted material to ensure correct in- the drawings of the male genitalia. terpretation of the state of pruinosity. Ratios and some structures discussed in this We use the term 'strong bristles' in describing study are abbreviated as follows: armature of the legs. Among ceratopogonids, L: length

Table 6. Descriptive statistics for hind femur length/width of male Serromyia.

Species n Mean Min. Max. SD barberi 10 6.51 5.62 7.06 0.575 borealis 2 5.67 5.57 5.77 — crassifemorata 9 6.10 4.16 7.06 0.937 nudicolis 16 5.96 5.38 6.64 0.388 sierrensis 1 6.19 6.19 6.19 - vockerothi 3 5.90 5.74 6.05 — ledicola 15 6.51 5.87 7.36 0.445 atra 13 6.55 5.00 7.00 0.539 bicolor 3 7.64 7.52 7.76 — femorata 19 6.47 5.23 7.52 0.517 mangrovi 2 4.12 4.08 4.15 — morio 13 7.15 6.32 7.50 0.612 pacifica 1 6.00 6.00 6.00 — rufitarsis 10 6.68 5.90 7.06 0.367 subinermis 10 6.82 6.46 7.26 0.287 tecta 2 6.28 6.25 6.32 — Revision of Holarctic species of Serromyia Meigen 175

Kieffer (1913, 1919) noted that the fork of the communis Fabricius 1805 and C.palustris cubitus (as posticale) was markedly distal to the Meigen 1804 as members of Serromyia but there position of the crossvein r-m (as transversale) is, in our opinion, no evidence for this. Remm and that the wings were spotted. Combined (1988) placed C. communis in Ceratopogon with the presence of short, equal claws on the (considering that Fabricius' description of fore and mid legs and an elongate hind claw C. communis was not a new name but fol- (equal to the length of the fifth tarsomere), lowed Meigen's earlier 1804 description) and this description cannot apply to any known C.palustris in Dasyhelea Kieffer. The date of Serromyia, but does fit a general description of submission for the catalogue in Remm (1988) several species of Palaearctic Monohelea. We was the end of 1982 and Szadziewski (1986), therefore transfer the name to that genus as a after examining the type specimens, has shown new combination: Monohelea scirpi (Kieffer). that the Ceratopogon palustris is actually a Serromyia fuligipennis Clastrier has recently species of Forcipomyia. Havelka (1978) also been placed in a new genus Congohelea Wirth interpreted Ceratopogon candidatus Winnertz & Grogan and we agree that it was incorrectly 1852 as a member of Serromyia but considered placed in Serromyia. the name unavailable because of lack of use Havelka (1978) considered Ceratopogon (50-year rule). Remm (1988) correctly placed

Figs 12A-D. A aedeagus, B paramere, C male genitalia of Serromyia tecta; D, female genitalia of S.borealis.