Revision of European species of Siphona Meigen (Diptera: Tachinidae)

Revision of European species of Siphona Meigen (Diptera:

Tachinidae)

STIG ANDERSEN

A revision of the European species of Siphona Meigen (1803) is presented. New diagnostic characters, especially based on structural differences in the male and female genitalia, are introduced. Also the traditionally used characters are revised. Four species are described as new: ingerae, mesnili and variutu from Denmark, and martini from Sweden. Thirteen other species are diagnosed. All seventeen species are keyed, and details of the genitalia are illustrated. Two lectotypes are designated. Notes on the affinities and distribution of the species are given. The biology is reviewed and possible host-parasite relationships are presented in an annotated list. True aerial swarming (synorchesia) of male Tachinidae is reported for the first time.

S. Andersen, Zool. Mus., Universitetsparken 15, DK-2100 Copenhagen, Denmark.

Contents

I Introduction ............................. 149 II Material and methods ..................... 150 III Inter- and intraspecific variation in characters 150

A. Size and other measurements ........... 150 B. Colour and pollinosity ............... I .151 C. Chaetotaxy ........................... 153 D. Abdomen ............................ 154 E. Male genitalia ......................... 154 F. Female genitalia ....................... 154

IV Bionomics ............................... 157 A. Host-parasite relations ................. 157 B. Feeding habits ........................ 158 C. Habitat preferences .................... 158 D. Swarming behaviour ................... 158

V Taxonomy ............................... 159 A. Diagnosis ............................ 159 B. Key to European species of Siphonu ..... 159 D. Treatment of the species ............... 161

VI Literature ............................... 171

I. Introduction

Flies belonging to the genus Siphona are easily recognizable by the long and slender geniculate proboscis with haustellum and labellum sclerotized, folded back in resting position, and with very specialized labellar structure. A long and slender proboscis is also found in many other Diptera, which in the past has caused con-

fusion in the proper application of the name Sip- hona, which has been assigned to both Cono- pidae and Muscidae. In his original diagnosis Meigen (1803) selected Stomoxys irritans Fabri- cius, 1775 (nec Linnaeus, 1758) (also with a long proboscis) as the type of his genus. Some of the characters he observed (arista bare and proboscis geniculate) showed that he probably had geniculata DeGeer or a closely allied species before him. Opinion No. 1008 of the Interna- tional Commission on Zoological Nomenclature (ICZN) has fixed the type-species of Siphona as geniculata DeGeer, with irritans Linnaeus, 1758 as the type-species of the muscid genus Haema- tobia, confirming an earlier designation.

The genus Siphona contains numerous externally rather uniform species occurring all over the World, except in Australia and New Zealand. In East Africa and Europe the genus is richly represented and many species have been described. Mesnil increased the number of Euro- pean species from 4 to 13, as listed in his com-

prehensive work on the Palaearctic Siphonini (1962-65). Based on external characters, Mesnil gives a very thorough description of all species, but many of his primary diagnostic features are not quite reliable, being trivial or unstable. The present revisionary work is restricted to Euro-

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pean species and is based mainly on new diagnostic characters, especially structural differences in the genitalia, but also traditionally used characters are revised. 4 new species have been discovered, which are fully described; for the species previously described only a diagnosis is presented here. Notes on types, distribution and biology and a key for identification of the species, are included.

The possible monophyly, and phylogenetic position of Siphona within the tribe Siphonini, was treated by Andersen (in press).

II. Material and methods

This study was primarily based on about 1800

specimens housed in the collection of the Zoological Museum of Copenhagen. Additional Swedish material was borrowed from the Museum of Zoology in Lund, collected mainly by Dr. H. Andersson. Primary types have been examined of 7 out of 16 species of which type material is known to be still in existence. For the remaining species type examination was not essential, because the species were sufficiently described by Dr. L. P. Mesnil or recently examined by Dr. B. Herting, who kindly sent me information on the types. Some specimens were also kindly sent to me by Dr. Herting (SMN) for dissection (S. grandistyla Pandellé (d'), silvarum

Herting and hokkaidensis Mesnil). Only the type material of S. genicu/ata (DeGeer) is assumed lost.

Regarding methods, especially for treating the male genitalia, see Andersen (in press).

The following abbreviations for depositories are used:

CNC Canadian National Collection, Ottawa, Ca- nada.

HDE Hope Department of Entomology, Oxford, England.

MNHN Museum National d'Histoire Naturelle, Pa- ris, France.

MZ Museo Zoologico "La Specola", Florence, Italy.

SMN Staatliches Museum fur Naturkunde, Lud- wigsburg, West Germany.

UZI Universitetets Zoologiska Museum, Lund, Sweden.

ZI Zoological Institute, Academy of Sciences, Leningrad, USSR.

ZMUC Zoological Museum of the University, Copenhagen, Denmark.

III. Inter- and intraspecific variation in characters

Some of the characters traditionally used for

separating European Siphona have proved very stable, providing good key-characters in some species, while in other species they are unstable and unreliable. Apparently, geographical variation is also important. Local, aberrant populations are found in some species, and the possi- bility exists that these populations have moved on to a different host. It is conceivable that such a shift could be an early step in speciation, resulting in so-called host-races (Bush 1975). To rec- ognize normal specific variation a large number of specimens is necessary. The following dis- cussion outlines some of the characters useful in the taxonomy, at the specific level, of Siphona and indicates that some of the characters hit- herto used are of little taxonomic value.

A. Size and other measurements. The body length is generally of some constancy, but dwarf-specimens are often found either singly (in most species) or in abundance (in local populations of cnllini and gerciculata). S. pauciseta and variata, are small species of about 3 mm and ingerae, cristata andf7at,ifrons are large species of 5-6 mm. There is no significant sexual difference in size. Proportional measurements of the head, the standards of which are outlined in Fig. 1, are generally useful, but unfortunately much intraspecific variation occurs. The lengths of face and antenna relative to the length of frons and the width of parafacials relative to the maximal width of the palp provide good specific characters in males, but are of less significance in females. Antennal segment 3 is normally narrow in females, but more or less widened in males, which when compared to the maximal width of the front femur provides important information for species recognition. However, some variation is noted in the shape of segment 3 of the male antenna. In confusa, collini and maculata it is mostly subrectangular with straight innerside, but occasionally it has a curved innerside and thus appears larger and thinner. In ingerae and mesnili the situation is just the opposite. Such structural variation also occurs in other Siphona and is perhaps to a high degree an artifact rather than true variation. The width of the gena is usually about 0.2 x the height

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Fig. 1. Male head of S. ingerae n.sp., standards of measurements, a-length of face, b-length of frons, c-height of head, d-height of eye, e-width of gena, f-length of haustellum.

of the eye, but in ingerae 0.3-0.4 x and 0.4-0.5 x the height of the eye in males and females, re- spectively ; this is a good specific character, at least for females. There seems to be no difference in the width of the frons relative to the width of head, neither between the sexes nor the species. The haustellum is usually shorter than the height of the eye in maculata. In ingerae and mesnili the length of the haustellum is varying between the height of the eye and the height of the head, and is in paludosa about equal to the

height of the head; in the remaining species it is usually distinctly longer than the height of the head. The length of the labellum relative to the haustellum is not used in European species, but has been used in the separation of North Ameri- can species. The number of lateral pairs of pseudotracheae of the labellum is 4 in ingerae, maculata and mesnili, 3-4 in collini, flavifron,s and geniculata and 2-3 in confusa, and does not seem useful in classification. Some species have

slightly longer tarsal claws than others, but this is only a distinct and good taxonomic character in males of ingerae.

B. Colour and pollinosity. The colour of the fa- cial pollinosity provides no useful characters. Usually the pollinosity of the parafrontals is silvery white and rather dense except around setae on the parafrontalia, giving them a dark

appearance, but in some species it is yellowish. Only in cristata is the pollinosity more invari- able ; yellowish to golden and very dense. The colour of antennal segments 1 and 2 is important. In most species it is more or less fuscous yellow, but bright yellow inflavifrons, rossica, paludosa and in some geniculata. In variata, the colour is whitish yellow. Antennal segment 3 is usually quite dark, but in some species often yellow proximally (a very variable character, used by Mesnil in paludosa and flavifYOns). Thorax is densely greyish or yellowish pollinose with distinct dark vittae only in ingerae and maculata; however, faint indications of vittae are found in several species. Colour of femora is very useful. These are usually entirely yellow or almost yellow, but darkened apically in boreata, martini and paludosa, and almost entirely dark, at least on front femur in ingerae, collini, and geniculata. The colour (yellowish or blackish) of the tegula is a primary diagnostic character used by Mesnil to distinguish between species. However, many species frequently have red or more or less fuscous yellow tegulae and are therefore difficult to place. These uncertainties have not been completely eliminated in the present key. The colour of the abdomen is of great taxonomic value. The ground colour of the male abdomen is usually dark with a characteristicly small to large field of yellow ground colour confined to the lateral parts of Tl-4 (except in ingerae, some paludosa and many confusa, cnllini and geniculata which are entirely dark in ground colour, and flavifrons and cristata. which are almost or entirely yellow). The ground colour of the female abdomen is usually entirely dark, but occasionally with small lateral fields of yellow ground colour confined to Tl-2, and usually with larger yellow fields in flavifrons, cristata, boreata, silvarum, martini, rossica and paludosa. The abdomen is densely greyish or yellowish pollinose except for dark spots around setae and setulae ; these spots are usually small, but some-

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Fig. 2. Aedeagus of- A. S. ingerae n.sp., lateral view showing tooth-bearing plates of distiphallus. - B. S. intrudens (Cn.) (U.S.A.), lateral view showing contours of the proximal part of distiphallus. - C. S. maculdta StTg. in Zett., lateral view showing tooth-bearing plates partly enclosed by the membrane. - D. S. ingerae n.sp., anterior view showing asymmetry in the position of the teeth. A & B: photomicrographs, C & D: scanning electron micrographs.

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Figs. 3-5. Structure of male St5. - 3. S. ingerae n.sp. - 4. S. mesnili n.sp. - 5. S. geniculata (DeG.). - Fig. 6. Female genitalia of S..setosa Mesnil. - Figs. 7-10. Structure of female sternites. - 7. S. geniculata (DeG.), St8. - 8. S. paludosa Mesnil, St7.-9. S. boreata Mesnil, St6.-IO. S. cristata (Fabr.), St6. - Figs. 11-17. Structure of cerci and surstyli. - 11. S. ingerae n.sp. - 12. S. mesnili n.sp. - 13. S. confuse Mesnil. - 14. S. variata n.sp. - 15. S. ?variata n.sp. (Spain). - 16. S. rossica Mesnil. - 17. S. geniculata (DeG.).

what larger in rossica, and sometimes coalesced and forming marginal bands in silvarum. In most paludosa there is a more or less regular median non pollinose dark vitta. C. Chaetotaxy. Features of the setae of the head are generally not useful for identification. The setulae on the parafacialia are usually weak and confined to the upper third of the face, but in most geniculata and ro.ssica they extend to the middle. The tip of the palp is normally setulose, but bare in cristata and bare, or almost so, in confuse, variata, boreata and grandistyla. The thoracic chaetotaxy is of great value. The number (3 or 4) of postsutural dc setae is important and of high constancy in most species, but variable in some. Basically, cristata, flavifrons, setnsa and rossica have 4 dc, but the second seta

is often small or indistinct. A very stable character is the absence of a presutural ia seta in collini, sometimes also absent in confusa. Usually 1-3 pairs of setulae are present on the prosternum, but this is always bare in cristata, setosa and paludosa, and sometimes also in flavifrons and geniculata. The chaetotaxy of the legs is important for species recognition. The front femora are without anterior setae in all species except martini which has a preapical av seta. This would seem a good diagnostic character, al- though this seta is weak on one femora of one of the three known specimens. A probable primitive feature is the row of pv setae on the front femur in ingerae, many mesnili and a very few maculata; in all other species this row is incom- plete, and bears only two strong setae.

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Characteristic, but not a stable character, is the

presence on the front tibia of two pv setae in most rossica specimens, and some goniculata and silvarum. The missing preapical ad seta of the mid femur of ingerae and collini seems to be a stable character and is also only rarely found in mandata. One or two setulae on vein Rl seems to be a rare feature in rossica and setosa, but of more constancy in grandistyla. Abdominal chaetotaxy is very stable in some species and unstable in others. One pair of lateral marginal setae on T 1 +2 is well developed in most species, small in ingerae, and indistinct in mandata and collini; two pairs are present in many rossica and some geniculata. One pair of median marginal setae on T +2 is a very variable feature in_flavi- fi-ons, setosa, geniculata. silvarum and rossica; the remaining species are normally without median marginals. The number of setae on T3 is generally 4 (a lateral and a median pair of strong setae), but usually 6 strong marginals are present in setosa and rossica, and occasionally also in other species. In the latter case the two most lateral pairs are usually inserted closer to each other than to the median pair, but in a few sotosa the setae are inserted an almost equal distance

apart (this was used as a keycharacter by Mes- nil). Features of the setae on the abdominal sternites show little variation except with regard to the point of insertion of a seta on the lobes of the male St5. In some species this seta is close to the middle of the inner edge, in others closer to the incision than to the apex of the lobes. Var- iation of this feature rarely occurs.

D. Abdomen. The preabdomen is very uniform and offers no characters except in the lobes of the male St5, normally having a simple rounded

shape (Fig. 5), but of a characteristic shape in ingerae (Fig. 3) and mesnili (Fig. 4).

E. Male genitalia. Features of the hypandrium, pregonites, postgonites and epandrium are of no use in species recognition and very few differences are found in the profile of the fused cerci (cercal plate) and surstyli (Figs. 11-17). An

astonishing feature is found in the surstyli of variata in which a pair of small posteriorly-inwardly directed setae are present (Figs. 14, 15). Unfortunately this character is not quite stable, with the setae apparently indistinguishable from the ground setulae in one of the five males available. The most reliable characters are found in

the profile structures of the distiphallus (Figs. 18-38), especially variations in shape combined with characters of the proximal, well sclerotized part (a more or less spinulose plate, sometimes with distinctive outline only in cleared specimens, cf. Andersen, in press) and the distal semi-sclerotized part (weaker sclerotized part with teeth or spinules characteristically inserted along the outer edge on a more or less distinct plate, Fig. 2). It is possible to distinguish between two types of distiphalli: an assumed primitive type with the distal tooth-bearing plates more or less isolated giving the distal edge a serrate appearance (in ingerac, maculata, mesnili and cnllini) and another probably derived type with the distal tooth- or spinule-bearing plates very delicate (in the remaining species, except variata, with a varying degree of isolation between the delicate plates, Figs. 22-24). The general appearance of the distal dentition offers very useful characters, but some variation occurs. For example, the teeth may be inserted asymmetrically (Fig. 2D), or the tooth next to the posterior sclerite may be missing on one side as in many oonfitsn (Figs. 25, 26) and some variata (Figs. 22, 23), boreata, and grandistyla. In species with spinule-like teeth some of the smallest may be missing. In species with very distinct toothbearing plates the two most posterior appear connected with or derived from the posterior sclerite. The remaining plates connected with or forming part of a sclerite, may be interpreted as a fragmentary anterior sclerite (cf. Fig. 2A). The profile outline of the distiphallus proves to be rather constant within the species, however, minor intraspecific differences are noted in the shape of the distal part, which may appear more or less pointed. This is obviously due to varying extensions of the posterior sclerite into the distal membranous part. The posteri- or sclerite is usually well developed, but is re- duced or missing in rossica, some paludosa and geniculclta, and often with somewhat irregular outline of no value for classification. No characters have been found in the distal membranous part, which is without a sclerotized acrophallus. No essential characters are found in the basiphallus and stalk of the distiphallus.

F. Femule genitalia. Only a few specific characters are found in the female genitalia. Generally, differences in shape of the sternites

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Figs. 18-28. Aedeagus of- 18. S. ingerae n.sp. - 19. S. maculata Staeg. in Zett. - 20. S. collini Mesnil. - 21. S. rreesnili n.sp. - 22. S. variata n.sp. - 23. S. variata n.sp. - 24. S. ? variata n.sp. (Spain). - 25. S. confusa Mesnil. - 26. S. confusa Mesnil. - 27. S. cristata (Fabr.). - 28. S. setosa Mesnil. bph - basiphallus, d.p - distal part of distiphallus, dph - distiphallus, l.p.sc - left arm of posterior sclerite, p.p - proximal part of distiphallus, pro - projection of posterior sclerite, st - stalk of distiphallus, t-b. pl - tooth-bearing plate.

are too small to be of practical use; a few types are illustrated (Figs. 6-10). A character of the posterior apodeme of St7 is remarkable. Usually

this is well developed, but may also be membranous, and in rossica and paludosa it is apparently always very short and membranous. St8

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Figs. 29-38. Aedeagus of - 29. S. grandistyla Pand. - 30. S. bore ata Mesnil. - 31. S. flavifrons Stxg. in Zett. - 32. S. geniculata (DeG.). - 33. S. pauciseta Rondani. - 34. S. silvarum Herting. - 35. S. hokkaidensis Mesnil. - 36. S. martini n.sp. - 37. S. rossica Mesnil. - 38. S. paludosa Mesnil.

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has usually 1-3 pairs of outwardly directed setulae, but a higher number occurs in collini, flavifrons, geniculata, silvarum, rossica and paludosa (Fig. 7). The supra-anal plate is only found in mesnili and some confusa. Lingulae are not developed on the subanal (postgenital) plate.

IV. Bionomic.s

A. Host-parasite relations. Notes on the repro- ductive habits and immature stages are given by Andersen (in press). Fragments of the life-history and descriptions of immature stages are given by Bloeser (1914) for S. plusiae Coquillett, by Roubaud (1906) for S. cristata (Fabricius) (probably misidentified S. geniculata (DeGeer), cf. Herting 1960:63), and in more detail by Rennie & Sutherland (1920). The complete life-history is described by Alma (1974), for the economically important species S. geniculata (DeGeer). The life-history of geniculata is briefly reviewed here. The larvae mainly attack larvae of Tipula paludosa Meigen, T. maxima Poda and T. oleracea Linnaeus. Multiple parasitism was shown by Alma to be common. As many as 15 parasitic larvae were found in one larva of T. maxima. Rennie & Sutherland found two generations a year in Scotland. The larvae hibernate within their hosts. Pupation begins in April, and after a pupal period of about three weeks the imagines emerge during April or May. Larvae from this generation of flies are thought to attack full- grown Tipula larvae. A second generation of adult flies appears towards the end of July, and the larvae of this generation probably attack young Tipula larvae. In Denmark a third generation of adult flies appears in September and does not die out until the end of October. In Great Britain, the percentage of parasitism found by Alma in third- and fourth-instar larvae of the overwintering generation of Tipula maxima was as high as 15.2 and 34.1, in two subsequent years. Tipula paludosa and oleracea are fairly common in cultivated soil and are known to cause much damage to crops, including grass. Carter (1976: 120) stated that of the parasites found in tipulids only two viruses, some nema- todes and a Siphona sp. were usually lethal to their hosts. He therefore proposed that the search for a biological control agent against harmful tipulids should be concentrated on these

parasites. According to Wilkinson (1971) S.

geniculata was released, in 1968, in a biological control programme in British Columbia against T. paludosa, as the latter had done much damage there to pastures and hay meadows. Since then no recoveries have been made and Wilkinson recommended new attempts to be made to es- tablish this tachinid fly.

Scattered information only exists about host- parasite relations in other Siphona species. The unreliability of identifications of tachinid parasites has resulted in much uncertainty about host records. The host list below is mainly derived from data in Mesnil (1964-65) and Herting (1960). The hosts listed for cristata probably in- clude four hosts (names with questionmark) of

cOl{fusa which was earlier confused with cristata. Most other parasite data are attributed to Mesnil, Herting or myself. Lundbeck (1927: 476) recorded two specimens of flavifrons reared from Eupithecia succenturiatu (Linnaeus). I have seen the setosa specimens myself; the records for this species are the first ever published. The unknown host of flctvifrons is probably tipulid larvae.

Parasite-host list

S. maculata LEP., Noctuidae: Euxoa obeLisca (De- nis & Schiffermuller), Ochropleura candelisequa (Denis & Schiffermül1er).

S. collini LEP., Noctuidae: Euxoa temera Hubner.

S. confusa LEP., Cerapteryx graminis (Linnaeus), Noctuidae: Phlogophora meticuosa (Linnaeus) (Radisson & Masson, 1974: 109), Mythimna litorais (Curtis).

S. cristata LEP. Geometridae: Anticullix sparsata (Treitschke), Erannis defoliaria (Clerck). LEP., Sphingidae: Smorinthus ocellata (Linnaeus). LEP., Noctuidae: Euxoa tritici (Linnaeus), Mamestra brassicae (Linnaeus), Lacanobia suasa (Denis & Schiffermuller) (C. S. Larsen), L. oleracoa (Linnaeus), Hadena bicruris (Hufnagel) (C. S. Larsen), ? Mythimna ,ferrago (Fabricius), ? M. litoralis (Curtis), ? M. ohsolc ta (Hbbner), Antitype chi (Linnaeus), ? Phlogophora meticulosa (Linnaeus), Caradrina morpheus (Hufnagel).

S. setosa LEP., Geometridae: Eupithecia succonturiata (Linnaeus) (I. C. Niel- sen). LEP. Noctuidae: Allophyes oxyacantae (Linnaeus) (C. S. Larsen).

S. geniculata DIPT., Tipulidae: Tipula subnodicornis Zetterstedt, T. varipennis Meigen, T. lateralis Meigen, T. montium Egger, T. oleracea Linnaeus, T. paludosa

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Meigen, T. fulvipennis DeGeer, T. maxima Poda, T. vittata Meigen. (cf. Alma 1974 and Carter 1976). LEP., Noctuidac: Ceramica pisi (Linnaeus) (Ford 1973: 289).

S. silvarum DIPT., Tipulidae: Tipula irrorata Mac- quart.

B. Feeding habits. Adult Siphona feed from flowers, especially those of various C't)ti7positae (cf. Draber-Monko 1978). The long geniculate proboscis is obviously an adaptation to reach the nectar in the corollatubes. The relatively short proboscis in species with an exclusively spring occurrence (see below) seems correlated with the fact that these species do not usually feed on tubuliflorous flowers. For example, maculata is most commonly observed on flowers of Sali.r, Prunus and Ribes, and mesnili on Anemone nemorosa. A few female ingerae were collected on young Betula with abundant honeydew on the leaves. Seasonal differences in the occurrence of Siphona species is significant. S. rnaculata, ingerae and rnosnili are exclusively springspecies, while oristata and flavU¡'ons are latesummer species with only one generation a year. In con-

,Illsa and pallciseta there seems to be at least two, and in cnllini and geniculata three, generations a year.

C. Habitat pre.fáellccs. Adults are collected in many habitats. Most species in my material were collected in woods. S. geniclllata, pauciseta, and collini occur more commonly on waste ground or cultivated fields. S. <.oqfi13.a seems to be exclusively coastal in Denmark. The most productive method of collecting is usually by sweeping flowers, but in some species males are very seldom collected on flowers, but can be found in certain sheltered places, where hundreds of males may occasionally engage in swarming activities.

D. Swarrning behaviour. Aerial swarming (cf. McAlpine & Munroe 1968) in the strict sense (synorchesia) is here reported for male Siphona, This is the first record in the Tachinidae and is, therefore, described in some detail. Herting ( 1960: 29) reported two kinds of mating behaviour in Tachinidae, namely one with a few male individuals engaged in pursuit flights from resting places (prosynorchesia) and one with a syrphid-like hovering as found in Carcelia plebe- rula Mesnil (mnnoorchc?sia). According to Baer

(1920) males of Sorvillia ursina (Meigen) are usually found singly, but sometimes accumulate at certain mating places, where they make quick rhythmic vertical and horizontal dancing movements and hover like a syrphid (polyorchesia). The dancing ("bilden auf- und abstiegend Reihen in einem Tanze ... ahnlich der Musca choroa des Fabricius" of some small Phasiinae observed by Robineau-Desvoidy and quoted by Mesnif ( 1944: 12) probably is due to a misidentification

(Mesnil, pers. comm.). In Siphol111. aerial

swarming has been observed in several species, but is most characteristic of ingerae and mesnili. All observations were made in North East Zea- land in the close vicinity of Copenhagen. The first observation, an aggregation of about 30 males, was made at noon on a warm, sunny day in spring. They flew close together, in a silent spiral-like flight with rapid zig-zag movements of the body and with the legs stretched out behind. Occasionally they remained stationary in mid-air (hovering) with the legs stretched out perpendicularly to the body, and all the time making brief aerial interceptions. During the period of observation (about two hours) the swarm was relatively stationary and did not change much in size. The individuals only rested for a few moments between the flights which, compared to later observations, were of long dura- tion indicating a peak swarming activity. Since then a few other swarming sites of ingerae have been found, at which swarming activity has been observed year after year. The males seem to appear a week earlier than the females, entering the swarming sites at once, and continue to swarm for about 20 days with peaks of activity on warm, clear days. On such days swarming of il1!?erae was observed above an elder bush under a tall spruce in another wood. The swarming began late in the morning with individuals swarming around the tips of the elder branches. At noon small aggregations of 2-5 individuals were engaged in brief aerial swarming. In the afternoon about 25 individuals were seen swarming more constantly in 2-3 aggregations between the bush and the lower branches of the spruce. In the evening more than a hundred individuals, forming several small swarms, were seen swarming almost constantly. Later on the swarming aggregations flow higher and higher up into the spruce-tree, until they could no longer be observed in the twilight.

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Males of mesnili were also seen swarming in tall spruces. In a particular tree about 30o individuals formed several small swarms, some near the top of the tree and some near the ground. Brief aerial swarming was also observed in rnactilata and geniclllata, but never with more than 5 individuals per swarm. In c.ollini, confusa. cristutcz and flavifi-ons only individual branch-tip swarming was noted. Females were usually present near the ground at the swarming sites, but they were never captured in the swarms. On several occasions coupling pairs (females newly hatched) were seen on the ground under the swarms. Aerial coupling in ingerae is indicated by the long claws of the male, which are pre- sumably used to clasp onto the females.

Within Siphonini aerial swarming has only been discovered in Siphona, but individual branch-tip swarming was once observed in Ceromya nigrohalterata (Villeneuve). The anal-vein is distinct to the wing margin in Sip- honia but not in C'oromyu. All members of the Siphonini have rather broad anal angles. The functional significance of a strong anal lobe in connection with aerial swarming is strongly sug- gested by McAlpine & Munroe ( 1968), who also presented the theory that swarming and associated morphological adaptation are primitive characteristics of all Diptera and the direc- tion of evolution is from social synorchesic swarming to individual swarming behaviour. In the ground plan of the Siphonini the anal vein is probably distinct to the wing margin (only shortened in Actia, Ceromya and Goniocera). This character is very rarely found in other Tachinidae, and is judged by Hennig (1958: 680) to be secondarily evolved within the Tachinidae. However, in view of the associated primitive mating behaviour found in Siphona and probably in the other Siphonini with a long anal vein, it seems assignable to the ground plan of the Tachinidac (cf. McAlpine & Munroe 1968: 1169).

V. Taxonomy

Genus Siphona Meigen

Crocuta Meigen, 1800. Name suppressed by ICZN (Opinion 1678, 1963: 339).

Siphona Meigen, 1803: 281. Type-species: Musca geniculata DeGeer, 1776: 20 by designation of ICZN (Opinion 1008, 1974: 157).

Bucentes Latreille, 1809: 339. Type-species: Bucc ntes cinerea Latreille, 1809 = Musca geniculata DeGeer, 1776, by monotypy.

Diagnosis: Proboscis long and slender, with haustellum and labellum well sclerotized, folded back in resting position, and labellum very specialized with only 2-4 pairs of pseudotracheae ; lower sternopleural seta stronger than the anterior upper seta; wing with anal vein reaching margin. d genitalia: Basiphallus without an epiphallus; distiphallus without distinct anterior sclerite, except in some species obviously with partitioned anterior sclerite present as tooth-bearing sclerites; posterior sclerite forked with usually short median projection, rarely with long projection; praegonite of normal triangular shape, bare. Q- genitalia : Ovipositor short with indistinct T6-7, St6-7 7 not bulging and with relatively long setae, and St7 usually with well developed apodeme. Parasites of various Macrolepidoptera and Diptera (Tipulidae). Occurring in almost all parts of the World, except Australia and New Zealand; 81 species.

Key to European species of Siphona

(Note. The present key is based mainly on external characters. Genital characters are in- dispensable in achieving an exact determination of atypical specimens or species closely resem-

bling one another).

1. Male: claws as long as last tarsal segment; female: gena about half as wide as eye height; abdominal ground colour entirely dark; mid femur without an ad preapical seta ...........

............................. 1. ingerae, n.sp. - Male: claws up to half as long as last tarsal

segment; female: gena up to one third as wide as eye height ................................. 2

2. T1+2 without strong marginal setae .......... 3 - T I + with distinct lateral marginal setae ...... 4 3. Strong presutural ia setae present; haustellum

usually shorter than eye height ............... ............. 2. maculata Stxger in Zetterstedt

- Strong presutural ia setae absent; haustellum longer than eye height ......... 3. collini Mesnil

4. Length of haustellum varying between eye height and head height. 3 postsutural dc setae; femora entirely yellow, front femur usually with more than 3 pv setae; antennal segments I and 2 fuscous yellow ................ 4. mesnili n.sp.

- Haustellum as long as or longer than height of head. 3 or 4 postsutural dc setae; front femur usually with 3 pv setae ...................... 5

5. Tegula yellowish or red, rarely fuscous red; width of male antenna never strongly exceeds maximal width of front femur ................ 6

- Tegula black or fuscous red; abdomen not extensively yellow ........................ 1 1

6. Lower half of parafacials in profile usually narrower than maximal width of palp; male face as well as antenna about 1.5 x as long as length of frons; segment 3 of male antenna subrectan-

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gular, about as wide as maximal width of front femur; femora entirely or mainly yellow; Tl+2 at most with a pair of median marginal setae of most varying size; parafacial setulae rarely de scending to middle of face ................... 7

- Lower half of parafacials in profile usually about as wide as or wider than maximal width of palp 8

7. Abdominal ground colour mainly mellow; antennal segments 1 and 2 yellow and not infuscated; T3 usually with 4 strong marginal setae; antennal segment 3 at least in female more or less yellow basally ..... 11. flavifrons Staeger in Zetterstedt

- Abdominal ground colour usually mainly dark; antennal segments I and 2 fuscous yellow; T3 with 5-6 strong marginal setae; prosternum bare ............................. 8. setosa Mesnil

8. T3-5 in male mainly with yellow ground colour; antennal segments I and 2 more or less fuscous yellow; face and antenna about as long as frons; prosternum bare; Tl 2 2 without median marginal setae; T3 with 4 strong marginal setae; femora entirely yellow; palp with tip bare or almost bare; parafrontals usually with dense yellowish to golden pollinosity ..... 7. cristata (Fabricius)

- T3-5 usually mainly with dark ground colour; prosternum with one pair of setulae (except in gc niculata with varying haircover ............ 9

9. 3 postsutural dc setae; face and antenna in male 1.3-2.Ox as long as frons, in female of about same length as frons; T1+2 without median marginal setae; female palp with tip bare or almost bare ............................. 10

- 4 postsutural dc setae; face and antenna usually up to 1. 3 x as long as frons; palp setulose or at least with some small setulae on the very apex; antennal segments 1 and 2 usually fuscous yellow ................................... 15

10. Antennal segments I and 2 more or less fuscous yellow; femora more or less fuscous yellow at least apically; distiphallus with the distal tooth- bearing plates not isolated from each other; surstyli with normal hair cover (Fig. 13) .......

........................... 6. confusa Mesnil - Antennal segments I and 2 yellow and not in-

fuscated ; femora entirely yellow; distiphallus with the distal tooth-bearing plates more or less isolated from each other; surstyli with isolated backward-inward directed setae (Fig. 14) ...... .............................. 5. variata n.sp.

11. Antennal segments 1 and 2 usually fuscous yellow; prosternum with one pair of setulae (except in geniculata with varying haircover); St7 of female with well developed apodeme (not known in grandistyla) ..................... 12

- Antennal segments 1 and 2 yellow and not or very slightly infuscated; face and antenna in male about 1.5 x as long as frons; lower half of parafacials in profile at most as broad as maximal width of palp in male, broader in female; abdomen usually with small lateral fields of yellow ground colour confined to Tl-3; St7 of female with small and weakly sclerotized apodeme (Fig. 8) .......................... 17

12. Antenna in male 1.5-2.0 x in female about 1.3x as long as frons; palp with tip bare or almost

bare; distal half of segment 3 of male antenna slightly broader than proximal half, and slightly broader than maximal width of front femur; if broader in the middle then distinctly broader than maximal width of front femur; T1+2 with out median marginal setae .................. 13

- Antenna in male rarely as much as 1.5 x as long as frons, in female about as long as frons; apex of palp more or less setulose; 4 postsutural dc setae

......................................... 14 13. Usually 4 postsutural dc setae present; lower

half of parafacials in profile narrower than maximal width of palp; mid and hind legs with fuscous knees; tegula black or very infuscated red; wings with Rl bare; female St6 heart-shaped (Fig. 9), St8 with 2-3 setulae laterally .........

........................... 9. boreata Mesnil - 3 postsutural dc setae; lower half of parafacials

in profile as wide as or wider than maximal width of palp, knees entirely yellow; tegula only slightly infuscated red; R 1 sometimes with 1-2 setulae apically ........ 10. grandistyla PandeII6

14. Antenna with segment 3 subrectangular, sometimes with very curved innerside and therefore widest in the middle, up to as wide as maximal width of front femur; lower half of parafacials in profile as wide as or wider than maximal width of palp ..................................... 15

- Distal half of segment 3 of male antenna distinctly wider than proximal half, and distinctly wider than maximal width of front femur; lower half of parafacials in profile narrower than maximal width of palp, with up to 10 setulae of normal size which are confined to upper third of face; Tl+2 usually without strong median mar ginals .................................... 16

15. T1 +2 with a pair of median marginal setae of varying strength, sometimes even missing; parafacials usually with more than 10 spinulose setulae which descend to or beyond middle of face; St8 of female setulose .................. ...................... 12. geniculata (DeGeer)

- T +2 2 without median marginal setae; parafacials with 0-6 fine setulae confined to upper third of face; palp at least with some setulae on the extreme tip; St8 of female with 1-2 pairs of setulae laterally ......... 13. pauciseta Rondani

16. Front femur without preapical av seta; female St8 setulose .............. 14. silvarum Herting

- Front femur with a preapical av seta; female St8 with one pair of setulae ....... 15. martini n.sp.

17. Distal half of segment 3 of male antenna distinctly wider than proximal half and distinctly wider than maximal width of front femur; T1+2 2 with a pair of median marginal setae, rarely without; parafacials usually with more than 10 spinulose setulae descending almost to middle of face, in some females even beyond middle; prosternum with 1-3 pairs of setulae; haustellum distinctly longer than height of head; front tibia usually with 2 pv setae; abdomen densely pollinose, except for large nonpollinose dark spots around setae; female St8 with 7-11 setulae ........................... 16. rossica Mesnil

- Distal half of segment 3 of male antenna not or

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slightly wider than proximal half, at most as wide as maximal width of front femur; TI +2 without median marginal setae; parafacials with at most 10 setulae of normal size, and confined to upper third of face; prosternum bare, rarely with a pair of minute setulae; haustellum as long as or slightly longer than height of head, front tibia usually with one pv seta; abdomen densely pollinose, except for small nonpollinose dark spots around the setae and a dark median vitta, which may be inconspicuous and only in posterior view indicated in the medio-dorsal region of T3; St8 with up to 7 setulae ....... 17. paludosa Mesnil

D. Treatment of'the species

1. Siphona ingerae n.sp. Figs. 1, 2AD, 3, 1 11 18.

Type locality: Denmark, NEZ, Bagsvaerd. Type material: Holotype d, Denmark: North East Zea- land, Bagsvaerd, Sm¢rmose, 26.IV.1979, S. Andersen (ZMUC). - Paratypes, 19 d with same data as holotype. - Further material seen but not considered as paratypes: Denmark: 15 d 3 Y, type-locality, 13.IV.-5. V. 1974-9, S. Andersen; 3 d 12, North East Zealand, Valby Park, 17.IV.1974, S. Andersen; 12, North East Zealand, Gevninge, 25.V.1977, S. Andersen; Funen, Bramstrup Mose, I.V. 1978, S. Andersen; Funen, 0rbask, Dyrehaven, LV.1978, S. Andersen; 6 d I 2, North East Zealand, Hareskov, 2.V.1978, S. Andersen; 1 d, South Jutland, Als, Birkep¢l, 13.IV.1979, V. Michelsen; 1 d, North East Jutland, Lerkenfeld, 18.IV.1979, S. Andersen. Diagnosis: Female gena 0.4-0.5 x as wide as height of eye; length of haustellum varying between height of eye and height of head; male claws as long as last tarsal segment (only about half as long as tarsal segment in other European Siphona); front femur fuscous yellow, with 3-5 pd and 4-6 pv setae; mid femur without a distinct preapical ad seta (elsewhere only found in collini) ; Tl+2 with a small but distinct pair of lateral marginal setae; d genitalia: St5 with very distinctive outline (Fig. 3).

Description Male

Heaci (Fig. 1): Parafrontals and parafacials whitish grey pollinose, pollinosity on parafrontals thin, resulting in a dark appearence. Anten- nal segments 1 and 2 fuscous yellow; face about 1.6x as long as length of frons; epistome rather protruding, gena 0.3-0.4x as high as an eye; parafacials broadly visible in profile, with maximal width (near the middle) at most slightly narrower than maximal width of palp, with about

6 setulae confined to upper fourth of face; antenna about 1.6x as long as length of frons; antennal segment 3 subrectangular and with more or less curved innerside, about as broad as, and when very broad in the middle, also conspicuously broader than maximal width of front femur; palp not or slightly clavate towards tip, very setulose, and with 2-3 setae below; length of haustellum varying between height of eye and height of head.

Thorax: With three broad, sometimes partly confluent, brown vittae; scutellum brown pollinose; 3 strong postsutural dc setae; presutural ia setae well developed; prosternum with one pair of lateral setulae.

Legs: Femora yellow except for some brown to blackish brown markings on most of the posterior and antero-dorsal surfaces of front femur and apically on the posterior surface of mid and hind femora; knees usually entirely yellow; claws as long as last tarsal segment; front femur with 3-5 pd and 4-6 pv setae, surface between these rows with some erect setae and setulae, anterior surface without setae; mid femur without preapical ad seta.

Wing.s: Tegula black or fuscous red, basicosta pale yellow.

Abdomen: Whitish grey pollinose, except for small dark spots around setae and setulae; ground colour entirely dark; an uncommon feature in other European Siphona. T1 +2 with one

pair of small lateral marginal setae, without median marginals; T3 with one median and one lateral pair of strong marginal setae; T3-5 with relatively large basal bands devoid of ground setulae, dorsally occupying a quarter to a third of respective tergites.

Genitalia (7 dissections): St5 as in Fig. 3, with apex of lobes slightly dilated and curved up- wards, and with a seta inserted near middle of the inner edge; cerci and surstyli as in Fig. 11, with the apical half of cerci very slender and slightly curved forward as seen in profile; distiphallus distinctive (Fig. 18, 2A), but very similar to that of maculata (Fig. 19, 2C), only differing in shape and extension of the wellsclerotized part. Both species have large teeth and tooth-bearing sclerite present as distinctly isolated plates with obscured proximal terminations; the most posterior teeth are more or less closely connected with the posterior sclerite.

Length of body: 5.5-6.0 mm.

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Female

Face about 1.4x as long as length of frons; gena 0.4-0.5 x as wide as height of eye; parafacials at most tapering below; antenna usually only slightly longer than length of frons; antennal segment 3 subrectangular with rather curved outline, at most as wide as maximal width of front femur; palp strongly clavate towards tip; femora with weaker or almost missing brown markings; claws about half as long as last tarsal segment.

Genitalia: St6-8 very similar to those of setosa (Fig. 6), St8 with 1-2 pairs of lateral setae.

Affinities. A very distinctive species easily recognized by the long claws in the male, the broad gena in the female, and the absence of a preapical ad seta on mid femur. It is probably most closely allied to two North American species, S. intrudens (Curran), and an undeter- mined species (seen by me), with which it agrees, for example, in having relatively long male claws, relatively wide female gena and a similar distiphallic structure. The two North American species both have a distinct preapical ad seta. Of European species ingerae seems most closely related to Inaculata, which agrees in having trivittate thorax, large antenna, short haustellum and very similar genitalia.

2. Siphona maculata Stager in Zetterstedt Figs. 2C, 19.

Siphona maculata StTger in Zetterstedt 1849: 3212; Lundbeck 1927: 473, Fig. 99 (Bucente.s); Mesnil 1965: 87 I , Taf. XXVI, Fig. 150.

Type material: The present species was first recognized as distinct by the Danish dipterist R. C. Stxger, who sent his description (in a manuscript that probably no longer exists, Siphonarum Daniœ) to Zet- terstedt, who published it, apparently without altera- tions, and without seeing the types himself. 27 d and 11 g located in ZMUC are labelled "Coll. Staeger" and are all regarded as original syntypes. I have labelled, and here designate as lectotype, a d labelled "Macu- lata Stag." and "d. Charl." by Stsger. The type locality is Charlottenlund, North East Zealand. One syntype with the labels "Maculatn Stxg." and " Q Charl." is not conspecific with the lectotype but is a Siphona geniculata (DeGeer). The remaining syntypes (now paralectotypes) are not labelled and are conspecific with the lectotype except for 2 Q of Siphona me.snili n.sp. Diagnosi.s: Width of female gena 0.2-0.35 of height of eye; haustellum usually shorter than height of eye, i.e. shorter than in any other European Siphona; 3 post-

sutural dc setac; presutural ia seta well developed; mid femur usually with a distinct preapical ad seta; Tl+2 2 without distinct marginal setae; d genitalia ( 12 dissections) : St5 almost as in gelliculata (Fig. 5), but with the strong seta inserted near the middle of the lobes; cerci and surstyli as in confi/Sa (Fig. 13) ; distiphallus rather distinctive (Fig. 19); y-genitalia (8 dissections): St6 with a rather distinctive heart-shaped outline, almost as in boreata (Fig. 9); St8 with 3 lateral setulae. Mutc rial examined: 128 d 90 y, from nearly all provinces of Denmark. Germany: I d, Potsdam. France: 1 d, Rambouillet. Sweden: 5 d 5 y, from Scania and Halland.

Distribution: A rather common species in North Europe. Also found in Central Europe. Records from Israel ( y, ssp. griseola Mesnil, 1970: 118) and Formosa (Hennig 1941, Ville- neuve det.) may be regarded as doubtful.

Affinities: More evidence is needed to clarify the taxonomic status of S. maculata griseola Mesnil (mentioned above), but it appears to be a distinct species closely related to maculata. Moreover, among European species ma(-lilata agrees most closely with S. ingerae n.sp.

3. Siphona collini Mesnil Fig. 20.

Siphona collini Mesnil, 1960: 188; Mesnil 1964: 858, Taf. XXVI, Fig. 151. Holotype d, Great Britain, Cam- bridgeshire, Burwell, 3.V.1937 (Coll. J. E. Collin, HDE).

Diagnosis: Length of haustellum normally exceeds height of eye; 3 postsutural dc setae; presutural ia seta absent, a feature not found in other European Siphona except rarely in confusa; mid femur usually without a preapical ad seta; Tl+2 without any strong marginal setae; d genitalia (8 dissections): outline of St5 almost as in geniculata (Fig. 5), but with the strong seta inserted near the middle of the lobes; cerci and surstyli similar to those of <onfiisa (Fig. 13); distiphallus as in Fig. 20, rather distinctive in having the distal tooth- bearing plates narrowly isolated from each other. Q- genitalia (6 dissections): St8 entirely covered with small setulae. Material examined. Denmark: 36 d 30 'j?, from East Jutland, North East Jutland and North Zealand. Nor- way : 8 d 25 Q, Opland. Sweden: 5 d 4 g, Scania and Smaland.

Distribution. Seems to be widespread and locally common throughout Europe. According to Mesnil (1964: 859) known from Belgium, France, Germany and USSR; also reported from Poland (Draber-Monko 1978), Austria (Herting 1973: 110), and Japan (Mesnil & Walcher 1968).

Affinities. A very distinctive species easily recognized by the missing presutural ia seta.

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This seta is normally present in all other Palae- arctic Siphona except S. efflatouni, described by Mesnil (1964: 864) on the basis of a single female from Egypt. S. collini seems most closely related to efflatouni, with which it also agrees in the absence of strong marginal setae on Tl+2. They differ in genal width, this being 0.2-0.3 times the height of eye as opposed to 0.4 times in effla- tinmi, and in proportional differences of the wing.

4. Siphona mesnili n.sp. Figs. 4, 12, 21.

Type locality: Denmark, NEZ, Gribskov. Type material: Holotype d, Denmark: North East Zea- land, Gribskov, Harager Hegn, 14. V.1979, S. Andersen (ZMUC). - Paratypes, 30 d 10 Q, with same data as holotype. Further material seen but not considered as paratypes: Denmark: 2 y (Coll. Staeger, see no . 2); 10 d 5 y, S¢nderborg, 27-30.IV.1891, W. Wijstnei; I d Zea- land, R. W. Schlick; 1 d, Geels Skov, 19.IV.1953, W. Buch; 8 d, Hareskov, 20.IV.1957, L. Lyneborg; 22 d 3 Q, type locality, 7.V.1978, S. Andersen; 1 d, North East Jutland, Bjerget ved Gj#1 , 31.V.1978, S. Andersen & V. Michelsen; 3 d 3 y, different localities on Born- holm, 25.V.1979, S. Andersen & V. Michelsen; Swe- den : I Q, Halsingborg, 21.V.1934, O. Ringdahl; 2 d 3 y, different localities in Scania, Halland and Smh- land, 14. V.-6.VI.1961-78, H. Andersson; Switzer- land : 2 d, Arosa, 24.VI.1955, 1900 m, O. Ringdahl (UZI). Diagnosis: Width of gena about 0.25 of height of eye; length of haustellum varying between height of eye and height of head; 3 postsutural dc setae; femora entirely yellow, front femur with 3 strong pd setae and 4-6 strong pv setae ; tegula fuscous-yellow to almost black; T1+2 with a pair of rather strong lateral marginal setae, but without median marginals-, d genitalia: St5 with very distinctive outline (Fig. 4); y genitalia: Supraanal plate distinct and divided into two small plates, each with 1-2 setulae (a similar structure is rarely found in confuse, but then the plate is without setulae).

Description

Male

Head. Parafrontals and parafacials greyish white pollinose, pollinosity on parafrontals thin, resulting in a dark appearance; antennal segments 1 and 2 fuscous ye'low; face about 1.5 x as long as length of frons; epistome not protruding far; gena about 0.25 x as high as an eye; lower half of parafacials in profile at most as wide as maximal width of palp, with 5-11 usually rather spinulose setulae confined to upper third of face;

antenna about 1.5 x as long as length of frons; antennal segment 3 subrectangular with more or less curved innerside, about as wide as, sometimes when very broad in the middle, conspicuously wider than maximal width of front femur; palp slightly clavate towards tip, above with a more or less regular row of setulae extending almost to the apex and also with 2-3 setae below; length of haustellum varying between height of eye and height of head.

Thorax: Distinct vittae absent; scutellum greyish pollinose; 3 postsutural dc setae; presutural ia seta well developed; prosternum with 1-2 pairs of lateral setae.

Legs: Femora entirely yellow; claws about half as long as last tarsal segment; front femur with 3 strong pd setae and 4-6 pv setae, posterior surface with some additional setae and erect ground setulae and anterior surface without setae; mid femur with a preapical ad seta.

Wings: Tegula fuscous yellow to almost black, basicosta yellow.

Abdomen: Yellowish grey pollinose, except for small dark spots around setae; lateral fields of yellow ground colour large, only separated by rather narrow median dark vitta, T4 more of less black and T5 entirely black in ground colour; T 1 +2 with one pair of rather strong marginal setae laterally, without median marginals; T3 with one median and one lateral pair of strong marginal setae; T3-5 with narrow basal bands devoid of ground setulae, dorsally occupying about one fifth of the respective tergites.

Genitalia ( 11 dissections): St5 as in Fig. 4, with apex of lobes abruptly curved inwards, making inner edge almost right-angled, and with a seta inserted near the middle ; cerci and surstyli as in Fig. 12, cerci distinctly clavate towards the tip; distiphallus as in Fig. 21, with rather distinctive structure: proximally broad, distally more or less tapering, teeth large and inserted on more or less distinctly isolated plates with obscured proximal terminations and with the two most posteri- or ones more or less closely connected with the posterior sclerite.

Longth of body: 4.5-5.0 mm.

Female

Face about 1.2x as long as length of frons; antenna slightly longer than length of frons; antennal segment 3 subrectangular with rather

164

curved outline, narrower than the maximal width of front femur; palp strongly clavate towards tip, with tip bare or almost bare; abdomen entirely dark in ground colour.

Genitalia (6 dissections): St6-8 very similar to those of setosa (Fig. 6), St8 with 2-3 pairs of lateral setulae; with distinct supraanal plate consisting of two small plates, each with 1-2 setulae.

Affinities: This species is very similar to con- YM?, but has distinctive genitalia in both sexes. The number of pv setae of front femur is usually, but not always, distinctive. The structure of the distiphallus seems intermediate in regard to the degree of isolation of the tooth-bearing sclerites between the condition found in ingerae/maculata (with well isolated sclerites), and collinilva- riata (with slightly isolated sclerites).

5. Siphona variata n.sp. Figs. 14, 22, 23.

Type locality: Denmark, NEJ, Frederikshavn. Type material: Holotype d, Denmark, Frederikshavn, VII.1881, H. J. Hansen (ZMUC). - Paratypes: Den- mark : 2 d, presumed same data as the holotype; I d, Saeby, VII.1881, H. J. Hansen, 1 d 2 y, North East Zealand, Melby Overdrev, 24.VI.1974, S. Andersen. Diagnosis: Closely resembling confusa, with following points of difference: antennal segments I and 2 bright yellow or very slightly infuscated yellow; femora entirely yellow or almost so; tegula yellow; d genitalia: surstyli usually with an isolated posteriorly and inwardly directed seta (Fig. 14); structure of distiphallus rather distinctive (Figs. 22, 23), with the distal tooth- bearing plates more or less isolated from each other; y genitalia: not, with certainty, distinguishable from those of confusa.

Description

Male

Head: Parafrontals and parafacials with dense greyish or yellowish white pollinosity. Antennal segments 1 and 2 bright yellow or very slightly infuscated yellow; face about 1.5x as long as frons; epistome slightly protruding; gena 0.25 x as wide as height of eye; parafacials with lower half in profile as wide as or wider than the maximal width of palp, with 3-6 fine setulae on upper third; antennae about 1.5 x as long as length of frons; antennal segment 3 subrectangular with

more or less curved innerside and width at the middle about as broad as or slightly broader than maximal width of front femur; palp clavate towards tip, with 2 setae below and a few setulae dispersed above; tip bare or almost bare; haustellum slightly longer than height of head.

Thorax: Without distinct vittae; scutellum greyish white pollinose; 3 strong postsutural dc setae; presutural ia seta well developed; prosternum with one pair of lateral setulae.

Legs: Femora entirely bright yellow, sometimes very slightly infuscated on the apical part of hind femur; claws half as long as the last tarsal segment; front femur with up to 3 strong pd and pv setae; mid femur with a preapical ad seta.

Wings: Tegula bright yellow or reddish yellow, basicosta bright yellow.

Abdomen: yellowish grey pollinose, except for dark spots around setae; large lateral fields of yellow ground colour, only separated by a rather narrow median, dark vitta, posterior half of T4 and T5 entirely black in ground colour; Tl +2 with one pair of strong lateral marginal setae, without median marginals; T3 with one median and one lateral pair of strong marginal setae; T3-5 with narrow basal bands devoid of ground setulae.

Genitalia (5 dissections). St5 similar to that of geniculata (Fig. 5); cerci and surstyli as in Fig. 14, with an outline similar to confusa (Fig. 13), surstyli with a distinctive isolated backwardly and inwardly directed seta in 4 of the 5 males available; structure of distiphallus similar to that of confusu (Figs. 25, 26), differs in having the distal tooth-bearing plates distinguishable and more or less isolated from each other (Figs. 22, 23).

Length of body: 3.0-3.5 mm.

Female

Antenna about as long as length of frons, antennal segment 3 narrower than maximal width of front femur; palp strongly clavate towards tip; abdomen entirely dark in ground colour.

Genitalia (2 dissections): St6-8 very similar to those of setosa (Fig. 6), St8 with 1-2 lateral pairs of small setulae; sub-anal plate absent.

Affinities: This species is very similar to confusa. The female is difficult to separate from

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female confusa which rarely may have antennal segments 1-2 only slightly infuscated and which more commonly occurs with only slightly darkened legs.

In addition to the specimens listed above I have seen I d 8 y from Spain, Almeria, 5-22.IV.1966, L. Lyneborg. They probably rep- resent the present species, but differ from Danish material as follows: generally larger, about 4 mm long; distal tooth-bearing plates of distiphallus not distinguishable, distal teeth somewhat irregularly inserted (especially on the one side, Fig. 24), both surstyli possess one inwardly and slightly backwardly directed seta, which is stronger on the left surstylus (Fig. 15). Moreover, the male has fuscous yellow antennal segments 1 and 2, but bright yellow femora. All other specimens have antennal segments and femora bright yellow and, therefore, closely re- semble the type-series.

6. Siphona confusa Mesnil Figs. 13, 26, 26.

Siphona confusa Mesnil, 1961 : 201; Mesnil 1964: 859, Taf. XXVII, Fig. 156.

Type-material: Holotype d, Sweden, Småland, Granna, 22.V.1956, H. Pschorn-Walcher (Coll. Mesnil, CNC, Ottawa) [not seen by myself, but examined by B. Herting, who kindly gave me information about the holotype, see below]. Diagnosis: Male: Antennal segments 1 and 2 fuscous yellow; face 1.3-2.Ox as long as length of frons; lower half of parafacials in profile, at least below, wider than maximal width of palp, with 5-9 setulae of normal size confined to upper third; antenna 1.3-2.Ox as long as frons; antennal segment 3 subrectangular, sometimes with very curved innerside and, therefore, very wide in the middle, wider than the maximal width of front femur; palp slightly clavate towards tip; haustellum normally longer than height of head; 3 postsutural dc setae (specimens with 4 dc setae extremely rare); prosternum with a pair of setulae; femora fuscous yellow at least apically, front femur with up to 3 strong pv setae; tegula more or less infuscated yellow; abdomen usually with large lateral fields of yellow ground colour, only separated by rather narrow median dark vitta; posterior half of T4 and T5 entirely black in ground colour, sometimes abdomen entirely dark, or nearly so, in ground colour; T1 +2 with a pair of strong lateral marginal setae, without median marginals; T3 with a median and a lateral pair of strong marginal setae; genitalia (31 dissections): St5 of normal appearance similar to that of geniculata (Fig. 5); cerci and surstyli as in Fig. 13; distiphallus rather distinctive (Figs. 25, 26) with the distal tooth-bearing plates not isolated from each other (not distinguishable).

Female: Antennae about as long as frons or only slightly longer; palp strongly clavate towards tip; tip bare or almost bare; abdomen entirely dark in ground colour; genitalia (15 dissections): St8 with 1-2 pairs of lateral setulae; subanal plate, at most, indistinct, without setulae. Material examined: Denmark: 20 d 24 East Jutland, West Jutland, North East Jutland, North Zealand and Bornholm. Greece: 2 d, Kalimnos and Kos. Israel: I d, Jaffa. Italy: I Q, Sicilia. Norway: 5 Q, Opland. Spain: 2 d 3 Y, Almeria and Granada. Sweden: 8 d 16 Scania, Halland and Blekinge.

Distribution: Widespread, but local, species throughout Eruope. Also known from Israel and ? Mongolia (2 d ? cnnfusa recorded by Herting 1968: 54).

Notes on the holotype. Herting (in litt.) noted the following differences from the description of Mesnil: The face is 1.4x as long as the frons. Parafacials (actual width, not seen in profile) are, at a minimum 2.5 x as wide as the arista. Gena seen in profile as wide as 0.2x the maximal di- ameter of an eye (in the female 0.25). Haustellum is a little longer than, or almost equal to, the height of head (never shorter).

Affinities : Externally this species is very similar to variata n.sp. and mesnili n.sp. and certain female specimens may be difficult to separate from variata, lacking distinctive characters of the genitalia.

7. Siphona cristata (Fabricius) Figs. 10, 27.

Stomoxys cristata Fabricius, 1805: 281; Lundbeck 1927: 471, Fig. 98 (Bucente.s); Mesnil 1964: 861, Taf. XXVII, Fig. 158 (Siphona). Holotype Denmark, Zealand (ZMUC) [examined]. Siphona c-hetoliga Rondani, 1865: 194. Holotype Y, Italy (MZ) [examined by B. Herting (1965: 194)]. Siphona palpina Zetterstedt, 1859: 6064. Holotype Sweden, Lindholmen (UZI) [examined].

Diagnosis: Male: Parafrontals usually with dense yellowish to golden pollinosity; antennal segments 1 and 2 more or less fuscous yellow; face slightly longer than frons; lower half of parafacials in profile as wide as, or wider than, the maximal width of palp, with 6-10 setulae on upper third; antenna about as long as length of frons, segment 3 subrectangular, sometimes with more or less curved innerside and therefore widest at the middle, rarely as wide as the maximal width of front femur; palp with bare tip; haustellum longer than height of head; 4 postsutural dc setae, but with the second one, in most specimens being small or indistinct and, therefore, apparently only with 3 dc setae;

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prosternum bare; femora entirely yellow; tegula more or less fuscous yellow, rarely blackish, abdomen with extremely large lateral fields of yellow ground colour, only separated by median dark vitta, very narrow on T1+2 and T3, widened on T4, and T5 entirely black. TI+2 with a pair of strong lateral marginal setae, usually without median marginals; T3 with a median and/or rarely two strong lateral pairs of marginal setae; genitalia (1 1 dissections): St5, cerci and surstyli similar to those of genic-iilata (Figs. 5, 17); distiphallus as in Fig. 27, with rather distinctive structure. Female: Antenna slightly smaller with more curved outline; abdomen with smaller lateral fields of yellow ground colour, at most confined to T l +2; genitalia (5 dissections): St6 with rather distinctive outline (Fig. 10); St8 with 1-2 lateral pairs of setulae. Material examined: Holotype 2, Denmark, Zealand. Denmark: 43 d 15 Y, from most provinces. France: 2 d, Rambouillet. Sweden: 2 d, Scania.

Distribution. Not common, but a widely distributed species in Europe. Mesnil (1964: 862) recorded one female specimen from Japan, but the species is not included in the list of Tachi- nidae from Japan (Mesnil & Walcher 1968).

8. Siphona setosa Mesnil Figs. 6, 28.

Siphona setosa Mesnil, 1960: I 91; Mesnil 1965: 876.

Typc material: Holotype d, USSR, Tolmatschevo, Luga, 12. VII. 1937, A. A. Stackelberg (ZI). Diagnosis: Male: Closely resembling flavifrons, with following points of difference: antennal segments 1 and 2 fuscous yellow; prosternum usually bare (with or without'setulae in_flai,ifrons ); tegula more or less fuscous yellow; abdomen with lateral fields of yellow ground colour of normal size, separated by a median dark vitta of varying width, posterior half of T4 and T5 entirely black; T 1 +2 usually with strong median marginal setae; T3 with 5-6 strong marginal setae (either with a median and one or two lateral pairs, or with the setae equally arranged); genitalia (8 dissections): St5 almost as in geniculata (Fig. 5), but with seta inserted closer to the middle; cerci and surstyli similar to those of confusa (Fig. 13); distiphallus with very distinctive structure (Fig. 28). Female: Differs from the male as follows: face about 1.3 x as long as frons; parafacials wider, but as in the male narrower than maximal width of palp, the tip of which is strongly clavate (slightly so in the male and in 2 jlavifrons); antenna about 1.3x as long as frons, segment 3 much narrower than maximal width of front femur; palp with tip almost bare; abdomen with small lateral fields of yellow ground colour, at most, confined to Tri+2, abdomen sometimes entirely dark; genitalia (6 dissections): As in Fig. 6: St6 with subrectangular outline; St7 with well developed apodeme; St8 with 1-2 pairs of lateral setae. Material examined: Denmark: 6 d I1 2, North and Middle Jutland, North East Zealand and Funen. Nor- way : 6 Q, Opland. Sweden: 8 d 14 Q, Scania, Halland, Smaland, Ndrke and Norrbotten.

Distribution. Scarce and local species in North and Central Europe. According to Herting (1962: 81 ), Mesnil (1965 : 877), Elsasser (1977: 66), and Draber-Monko (1978: 181) earlier recorded in a few specimens from Austria, Germany, ? Great Britain, Poland, Sweden and USSR.

9. Siphona boreata Mesnil Figs. 9, 30.

Siphona borc ata Mesnil, 1960: 190; Mesnil 1964: 856, Taf. XXVI, Fig. 154.

Type material: Holotype d, West Germany, Arnsberg, 15.VI.1956, B. Herting (Coll. Mesnil, CNC, Ottawa). Diagnosis: Male: Antennal segments 1 and 2 fuscous yellow; face 1.5-2.Ox as long as frons; lower half of parafacials in profile narrower than maximal width of palp, with 3-7 setulae on upper third of face; antenna 1.5-2.0 x as long as frons; apical half of antennal segment 3 slightly wider than basal half and slightly wider than maximal width of front femur; palp with tip bare or almost bare; haustellum usually longer than height of head; 4 postsutural dc setae, specimens with only 3 dc setac extremely rare ('? = grancli.styla); prosternum with one pair of setulae; knees of mid and hind legs more or less fuscous, tegula black or fuscous red; abdomen with large lateral fields of yellow ground colour, only separated by a narrow median dark vitta, posterior half of T4 and T5 entirely black in ground colour; TI+2 without median marginal setae, with a pair of laterals; T3 with a median and a lateral pair of strong marginal setae; genitalia (6 dissections): St5 as in geniculata (Fig. 5); cerci and surstyli as in confusa (Fig. 13); distiphallus with distinctive structure (Fig. 30), but not definitely distinct from that of boreata (Fig. 29). Female. Face about 1.5 x as long as frons; antenna about 1.3x as long as frons; antennal segment 3 narrower than maximal width of front femur; abdomen with the small lateral fields of yellow ground colour, usually confined to Tl+2 and lateral parts of T3; genitalia (4 dissections): St6 with rather distinctive heart-shaped outline (Fig. 9), elsewhere only found in maculata; St8 laterally with 2-3 pairs of setulae. Material exnrnincd: Denmark: I d I Lolland and North East Zealand. Finland: 3 d 2 Q, Tavastia au- stralis and Karelia borealis. Sweden: 3 d 4 Q, Scania, Halland, Varmland and Lappland.

Distribution. A very rare North and Central

European species. In addition to the Scandina- vian material, 13 specimens are known from Austria, Germany, Great Britain, and USSR (Mesnil 1964: 858; Herting 1973: 110).

Notes on the type-material: The male holotype seems, according to Mesnil's description, typical of the species in most details, but the face is unusually short, only 1.2 x as long as frons. Also the female does not seem quite typical, as frons

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and face are of equal length. All Scandinavian female specimens have both the face and antenna distinctly longer than the frons.

Affinities : This species is very similar to grandistyla. Moreover, boreata is similar too, and in Mesnil's key (1964: 854) not reliably separated from paucisota (delicatula Mesnil) and silvarurrc (hokkaidensis Mesnil), but can be easily distinguished by the combination of characters listed in the diagnosis.

10. Siphona grandistyla PandeII6 Fig. 29.

Siphona grandistyla Pandellé, 1849: 108; Mesnil 1965: 868; Herting 1967: 9. Holotype d, France, The Pyrenees (MNHN).

Diagnosis: Male. Closely resembling boreata, with following points of difference: lower half of parafacials in profile about as wide as maximal width of palp; face and antennae shorter: 1.2x (in holotype according to Mesnil 1965: 868-9) and 1.5x (in my material) as long as frons; antenna] segment 3 subrectangular, with curved innerside and therefore widest in the middle, distinctly wider than maximal width of front femur; only 3 strong postsutural dc setae; femora almost entirely yellow, only hind femur slightly infuscated apically ; tegula infuscated red; wings with 1-2 setulae inserted apically on Rl, but sometimes without (Her- ting in mitt.), a feature rarely found in setosa and rossica (holotype); genitalia (I dissection): St5, cerci and surstyli similar to those of boreata (Figs. 5, 13); distiphallus (Fig. 29) not definitely distinct from that of boreata (Fig. 30). Female. According to Herting (1967: 9) it differs from the male as follows (compared to Mesnil's description) : parafacials at their narrowest (actual width, not seen in profile) almost half as wide as antennal segment 3 (only about 1/3 as wide as in boreata); antennal segment 3 about 3 x as long as segment 2 (about 4 x as long as in boreata), and almost 3 x as long as wide; abdomen densely olive-grey pollinose, entirely dark in ground colour; genitalia not studied. Material examined: Switzerland: I d, Delémont, Do- mont, 27.VI.1969 (Coll. Herting). An additional male from Norway (Troms, Gratangen, 29.VI.1956, H. Andersson leg.) probably belongs to this species, only differing in having slightly darkened hind knees and bare R 1.

Distribution: Very rare. An additional male is recorded from the Pyrenees, Cauterets (Coll. Mesnil) and according to Herting (in litt.) only a few males and a single female are known from Switzerland, Del6mont (Coll. Herting) and I specimen from Austria (Herting 1971: 110).

Affinities: The differences between boreata and grandistyla, as listed in the diagnosis, seem

rather small and some of them may not be quite stable. However, in my opinion they are distinct species, but the variability of certain characters, including female genitalia, need to be clarified on the basis of more material.

II. Siphona flavifrons Stseger in Zetterstedt. Fig. 31.

Siphona ,flavifrons Stager in Zetterstedt 1849: 321 1; Lundbeck 1927: 475 (Bucentes); Mesnil 1965: 865, Taf. XXVII, Fig. 157 (Siphona). Type material : Similar to maculata (see no 2) this species was described by Stwger in a manuscript, Siphonarum Daniae, sent to Zetterstedt. 1 d 2 y located in ZMUC, Copenhagen are labelled "Coll. St*ger" and are considered syntypes. B. Herting has labelled, and I now designate as lectotype, a Y syntype labelled "Flavifron.s Stxg." and " Q Ordrup". The type-locality is Ordrup, North East Zealand. The remaining paralectotypes are without labels and are conspecific with the lectotype. Dia,qnosi.s: Male: Antennal segments I and 2 yellow and not infuscated; face about 1.5 x as long as frons; lower half of parafacials in profile narrower than maximal width of palp, with 6-1l setulae normally confined to upper third of face; antenna about 1.5 x as long as frons, segment 3 subrectangular, about as wide as maximal width of front femur; palp with setulose tip; haustellum longer than height of head; usually 4 postsutural dc setae, but the second seta sometimes small or indistinct and then only 3 strong setae appear; femora entirely yellow; tegula not or slightly infuscated yellow; abdomen with entirely yellow ground colour or almost so, occasionally separated by a very narrow median dark vitta and T5 with black ground colour; T 1 +2 with a pair of strong lateral and usually a pair of median marginal setae of extremely variable size, sometimes indistinct; T3 with a median and one or two lateral pairs of strong marginal setae; genitalia (15 dissections) : St5, cerci and surstyli very similar to geniculata (Figs. 5, 17); distiphallus with rather distinctive structure (Fig. 31), but also closely resembling geni- crrlata (Fig. 32). Female: Face about 1.3 x as long as frons; lower half of parafacials about as wide as maximal width of palp; antenna usually slightly longer than frons, segment 3 much narrower than maximal width of front femur; ground colour of abdomen similar to that of the male, but more commonly with the posterior parts dark; genitalia (6 dissections): closely resemblinggeniculata. Material examined : Austria: 1 c¡?, Schwarzau. Den- mark : 29 d' 26 y, from most provinces. Norway: 2 d 6 Q, Opland and Rogaland. Sweden: 38 d 28 Q, from most provinces. Switzerland: 2 d Wallis, alt. 2000 m. West Germany: 2 d 1 Q, Schwarzwald.

Distribution. Widespread and locally common species in North and Central Europe. Towards the north, in Sweden, reaching Lule Lappmark and Norrbotten.

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12. Siphona geniculata (DeGeer) Figs. 5, 7, 17, 32. Musca geniculata DeGeer, 1776: 20, Figs. 19-23; Lundbeck 1927: 474, Figs. 100-101 (Bucentes); Mesnil 1965: 866, Taf. XXVII, Fig. 159 (Siphona). Type(s) ? sex, Europe (lost). Stomoxys minuta Fabricius, 1805: 282. Holotype <j', Denmark (ZMUC, Copenhagen) [examined]. Siphona analis Meigen, 1824: 157. Holotype d (MNHN, Paris) [examined by Herting 1972: 3]. Siphona cinerea Meigen, 1824: 156. Holotype $(MNHN, Paris) [examined by Herting 1972: 4]. Siphona nigrovittata Meigen, 1824: 157. Holotype d (MNHN, Paris) [examined by Herting 1972: 11 ]. Siphona tachinaria Meigen, 1824: 156. Holotype d, Germany (MNHN, Paris) [examined by Herting 1972: 13].

Diagnosis: Male: Antennal segments 1 and 2 more or less fuscous yellow, rarely yellow; face usually only slightly longer than frons; lower half of parafacials, in profile, as wide as or wider than maximal width of palp, with more than 10 spinulose setulae descending to or beyond middle of face; antenna usually slightly longer than frons; antennal segment 3 subrectangular with slightly curved innerside, rarely very wide at the middle and reaching the maximal width of front femur; palp with a variably setulose tip; haustellum usually longer than height of head; 4 postsutural dc setae; prosternum with a pair of setulae, sometimes bare; femora predominantly fuscous yellow; tegula more or less fuscous yellow or red, sometimes blackish; abdomen at most with large lateral fields of yellow ground colour, only separated by a rather narrow median dark vitta, posterior half or more of T4 and T5 entirely black, less often with a narrow median vitta, or abdomen wholly dark; T1+2 with I or 2 pairs of strong lateral and usually a pair of median marginal setae of varying size; sometimes without the median pair of setae in Danish material, normally these setae absent in Finnish material, in 18 d 15 $ of ? geniculata from Mongolia (Herting 1973: 17), and in "subspecies pauciseta " Mesnil, 1964: 856; T3 with I or 2 pairs of strong lateral and a pair of strong median marginal setae; genitalia (23 dissections): St5 as in Fig. 5, with seta inserted closer to incision than to apex of the lobe; cerci and surstyli as in Fig. 17, with a characteristic dilation shortly before apex of cerci; structure of distiphallus (Fig. 32) rather distinctive, but very similar to that of flavifrons. Female: Face about as long as frons; antennal segment 3 much narrower than maximal width of front femur, palp usually distinctly clavate towards tip, abdomen densely pollinose, except for small dark nonpollinose spots around setae, normally entirely dark in ground colour; genitalia (9 dissections): St8 usually with more than 8 setulae as in Fig. 7. Material examined. Denmark: 580 d 9, from nearly all provinces. Finland: 30 d 5 Karelia borealis. France: 4 d 2 <j', Paris. Norway: Stavanger. Sweden: 20d 28 y, Scania, Halland and Smaland. West Germany: 5 d, Nurnberg.

Distribution. Very common species through-

out Europe. Also known from Mongolia (see above) and ? Canada (released, but no recoveries have as yet been made according to Wilkinson 1969: 55). Other records (Sabrosky & Arnaud 1951 ) from North America are assumed uncer- tain (cf. Cole 1969: 570).

13. Siphona pauciseta Rondani Fig. 33.

Siphona pauciseta Rondani, 1865: 193. Types ? sex, Italy (MZ, Florence) [examined by Herting (1969: 198)]. Siphona oculata Pandellé, 1894: 108. Lectotype d, France (MNHN, Paris) [designated and examined by Herting ( 1978a: 6)]. Siphona delicatula Mesnil, 1960: 190; Mesnil 1964: 862, Taf. XXVII, Fig. 155. Holotype d, Great Britain (HDE, Oxford) [synonymized with pauciseta by Her- ting ( 1969: 198) 1.

Diagnosis. Male: Antennal segments 1 and 2 fuscous yellow; face about 1.2 x as long as frons; lower half of parafacials in profile usually about as wide as maximal width of palp, with 0-6 fine setulae on upper third of face ; antenna at most slightly longer than frons, antennal segment 3 subrectangular, sometimes with curved innerside and then widest at the middle and reaching the maximal width of front femur; palp with several short setulae on apex; haustellum longer than height of head; 4 postsutural dc setae; prosternum with a pair of setulae; femora almost entirely yellow, only hind femur and sometimes mid femur slightly infuscated apically ; tegula more or less fuscous yellow, sometimes black; abdomen with large lateral fields of yellow ground colour, only separated by a narrow median dark vitta, posterior half of T4 and T5 entirely black; T 1 + with a pair of strong lateral marginal setae, without median marginals ; T3 with a median and a lateral pair of strong marginal setae; genitalia ( 15 dissections): St5, cerci and surstyli similar to those of geniculata; distiphallus with distinctive structure (Fig. 33). Female: Antennal segment 3 subrectangular, much narrower than maximal width of front femur; abdomen with lateral fields of yellow ground colour at most confined to T1+2; genitalia (6 dissections): St8 with 1-2 pairs of lateral setulae.

Material examined: Denmark: 25 d 24 g, from most provinces. Sweden: 7d, Scania and Halland.

Distribution: Seems to be a local but widespread species throughout Europe. Besides the type material and my own material, only a few specimens are recorded from Austria, Great Bri- tain and USSR (Mesnil 1964: 864; Elsasser 1977: 66) and Yugoslavia (Herting 1978b: 4).

14. Siphona silvarum Herting. Fig. 34.

Siphona silvarum Herting, 1967: 9; Mesnil 1965: 869,

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Taf. XXVI, Fig. 153 (d S. hokkaidensis Mesnil). Holo- type d, West Germany, Freiburg im Breisgau, Moos- wald, reared from Tipula irrorata Meigen by Mesnil (Coll. Mesnil, CNC, Ottawa).

Diagnosis: Male: Resembling geniculata, with following differences: face about 1.3 x as long as frons; lower half of parafacials slender in profile, distinctly narrower than maximal width of palp, and with 6-9 setulae of normal strength confined to upper third of face; antenna about 1.2x as long as frons; apical half of antennal segment 3 conspicuously wider than basal half, and conspicuously wider than maximal width of front femur; femora almost entirely yellow, only hind femur sometimes infuscated apically; tegula black or blackish; abdomen with large fields of yellow ground colour; TI +2 with a pair of strong lateral and usually without strong median marginal setae [with a median pair of setae in some males in Coll. Mesnil (Mesnil 1975: 1400)]; genitalia (4 dissections: St5, cerci and surstyli similar to those of geniculata, but cerci not, or only slightly, dilated shortly before apex; distiphallus rather distinctive (Fig. 34), only closely resembling martini n.sp. (Fig. 36) and a Japanese species S. hokkaidensis Mesnil (Fig. 35), but apparently at least distinguishable from these two species by the more numerous distal teeth. Female. Differs from male as follows: Face and antenna only slightly longer than frons, segment 3 subrectangular, much narrower than maximal width of front femur; abdomen usually densely pollinose, except for dark spots around setae, but Scandinavian specimens may be rather sparsely pollinose especially around setae and marginal ground setulae, resulting in more or less conspicuous nonpollinose marginal bands; abdominal ground colour dark, except for small lateral fields of yellow, confined to the most lateral parts of TI-3; genitalia (3 dissections); St8 with more than 10 setulae as in genicr?lata (Fig. 7). Material examined. Finland: 1 d' Tavastia au- stralis and Karelia borealis. Sweden: 1 d' 2 g, Halland, SmAland and Varmland. Switzerland: I d Del6- mont. West Germany: 1 d', Stuttgart.

Distribution. A rare North and Central Euro- pean species. According to Mesnil (1965: 871) previously known from Austria, France, Germany, Switzerland (many specimens) and USSR.

Affinities. Very similar to martini n.sp., but easily separated by features of the front femur (see later). Moreover, silvarum very closely re- sembles a Japanese species S. hokkaidensis Mesnil (1957: 36), described from a single female, and was at first confused with this species (the holotype was described by Mesnil (1965 : 869) as the male of hokkuidensis), but later separated and described by Herting (1967: 9). Unfortunately, this separation was based on such unreliable characters as the presence or absence of median marginal setae on T1+2

(Mesnil 1975: 1400) and on features of the female abdominal pollinosity, which are found to be variable in Scandinavian silvarum (see above). I have examined 1 ç hokkaidensis (Mesnil det.), Japan, Sounkyo, Hokkaido (Type-locality), 20.IX.1961 and 1 d ?hokkaidensis, Japan, Hiko- san, Kyushu, 27. VII.1961, kindly sent to me by B. Herting. These two specimens seem to be conspecific and differ from silvarum in the following points: female abdomen with relatively large fields of yellow ground colour, only separated by a very narrow median dark vitta, and with a narrow posterior non-pollinose shining black marginal band on T3, T4-5 entirely black in ground colour; female palp only very slightly clavate towards tip; female St8 with only 5 setulae; male cerci more slender; distiphallus with rather distinctive structure (see above) (Fig. 35). In contrast to most other hokkaidensis, these specimens have no median marginal setae on T t +2 and only 3 dc setae in the male which is certainly aberrant. More material is needed to clarify the true taxonomic status of this rare and difficult species.

15. Siphona martini n.sp. Fig. 36.

Type locality: Sweden, Scania, Hyllstofta. Type material: Holotype d, Sweden: Scania, Hylls- tofta, 1-15. VII.1949, J. Agrell (UZI, Lund). - Para- types, Sweden: 1 d, Smaland, S. Unnarud, Ekhult, 25.VII.-8.VIII.1978, S. Andersen; Halland, Ens- löv, 16.VII. 1956, in sedge-moor, H. Andersson. Diagnosis: Closely resembling silvarum, with following differences: female palp with tip bare; front femur with a preapical av seta (not found in other European Siphona). d genitalia: Distiphallus (Fig. 36) only slightly different from that of silvarum (Fig. 34), but apparently distinguishable from this by fewer distal teeth. y genitalia: St8 with only one pair of lateral setulae.

Description

Male

Head: Parafrontals and parafacials with rather dense greyish or yellowish white pollinosity; antennal segments 1 and 2 fuscous yellow; face 1.3 x as long as frons; epistome slightly protruding ; gena 0.2 x height of eye; lower half of parafacials in profile distinctly narrower than maximal width of palp, and with setulae of normal strength confined to upper third of face; antenna about 1.3 x as long as frons; apical half of anten-

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nal segment 3 conspicuously wider than basal half and conspicuously wider than maximal width of front femur; palp slightly clavate towards tip, with 1-3 setae below and a few setulae dispersed above and apically; haustellum longer than height of head.

Thorax: Without vittae; scutellum yellowish grey pollinose; 4 postsutural dc setae, the two anterior ones of equal strength; presutural ia seta well developed; prosternum with a pair of lateral setulae. Legs: Femora yellow, except for the apices, which are very slightly infuscated on front femur, but more darkened on mid femur (type-material lacking hind legs); claws half as long as last tarsal segment; front femur normally with well developed av seta (weak on the one side in one specimen), with 2-3 strong pd and pv setae; mid femur with a preapical ad seta.

Wings: Tegula fuscous red, basicosta bright yellow.

Abdomen: Densely yellowish grey pollinose, except for small dark spots around setae; large lateral fields of yellow ground colour, only separated by a rather narrow median dark vitta, posterior half of T4 and T5 entirely black in ground colour; Tl+2 with a pair of strong lateral marginal setae, without median marginals; T3 with a lateral and a median pair of strong marginal setae; T3-5 with narrow basal bands devoid of ground setulae.

Genitalia (2 dissections): St5, cerci and surstyli very similar to those of geniculata (Figs. 5, 17), but with cerci more slender and not, or only slightly, dilated shortly before the apex; distiphallus of distinctive structure (Fig. 36), but very similar to silvarum (Fig. 34), differing significant- ly from the latter only by fewer distal teeth. Distinguishable from the Japanese species S. hokkaidensis Mesnil (Fig. 35), by slightly different outline; all three of these species possess small, spinule-like distal teeth inserted on a common, lightly sclerotized, plate-like sclerite.

Length of body: About 5.0 mm.

Female

Face and antenna only slightly longer than frons; antennal segment 3 subrectangular, narrower than maximal width of front femur; palp with tip bare; apical parts of mid and hind femora strongly infuscated; abdomen with lateral fields of yellow ground colour occupying most of

T1+2 and antero-lateral parts of T3, on T1+2 only separated by a narrow median dark vitta.

Genitalia ( 1 dissection): St6-8 very similar to those of .retosa (Fig. 6), but St8 with only one pair of lateral setae.

Affinities. A distinctive species easily recognized by the preapical of seta on front femur. Most closely resembling silvarum and the Japa- nese hokkaidensis. Moreover, the female is similar to the female of boreata (also with slender parafacials, bare tip of palp and relatively large fields of yellow ground colour of abdomen), but easily separated from this by the features of front femur, antenna and St6.

16. Siphona rossica Mesnil Figs. 16, 37.

Siphona rossica Mesnil, 1961: 202; Mesnil, 1965: 874. Holotype d, USSR, Leningrad, Lugsk, 8-11. VI.1925, A. A. Stackelberg (Zl).

Diagnosis. Male: Antennal segments I and 2 yellow or very slightly infuscated; face about 1.5 x as long as frons; lower half of parafacials in profile up to as wide as maximal width of palp, usually with more than 10 spinulose setulae at most descending to middle of face; antenna large, about 1.5 x as long as frons, apical half of segment 3 distinctly wider than basal half and distinctly wider than maximal width of front femur; palp with more or less setulose tip; haustellum distinctly longer than height of head; 4 postsutural dc setae, but the second seta, in most, small or indistinct and then only with 3 strong dc setae; prosternum with 1-3 lateral pairs of setulae; femora almost entirely yellow, only hind femur sometimes infuscated apically; front tibia usually with 2 pv setae; tegula black or blackish; abdomen usually with, sometimes without, small lateral fields of yellow ground colour confined to TI-3, and with large nonpollinose dark spots around setae; T1 +2 with 1-2 pairs of strong lateral and usually a pair of strong median marginal setae; T3 with 4-6 strong marginal setae; genitalia (8 dissections): outline of St5 similar to that of geniculata (Fig. 5); cerci and surstyli with rather distinctive outline (Fig. 16); distiphallus very distinctive (Fig. 37). Female: Face about 1.2 x as long as frons; lower half of parafacials in profile, at least above, wider than maximal width of palp and with more numerous setulae, descending to or beyond middle of face ; antenna about as long as frons, segment 3 subrectangular, narrower than maximal width of front femur; genitalia (4 dissections) : St7 similar to that of paludosa (Fig. 8), with short and lightly sclerotized apodeme; St8 with 7-11 1 setulae as in geniculata (Fig. 7). Material examined. Austria: 1 d I Y, Burgenland, Ilmitz. Denmark: 5 d, North East Zealand and North West Jutland. Spain: 3 d 3 Y, Granada. Sweden: 3 d I Q, Scania, Ostergotland and Dalarna.

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Distribution. A very rare but widely distributed species. Apart from the holotype, and my own material, only known from 1 d 1 y from USSR (Mesnil 1965: 876), 3 specimens from Au- stria (Herting 1973a: 110), and 1 d from Grmany (Herting 1978b: 5).

17. Siphona paludosa Mesnil Figs. 8, 38.

Siphona paludosa Mesnil, 1960: 188; Mesnil 1965: 872, Taf. XXVI, Fig. 152. Holotype d, USSR, Leningrad, Luga, Tolmatschevo, 22. VIII. 1935, A. A. Stackelberg (ZI).

Diagnosis: Male: Closely resembling rossica, with following differences: parafacials with up to 10 setulae of normal size at most confined to upper third of face; apical half of antennal segment 3 not or only slightly wider than basal half, up to as wide as maximal width of front femur; haustellum only as long as or slightly longer than height of head; prosternum bare, or rarely, with a pair of minute setulae; femora usually infuscated yellow apically; front tibia with a pv seta; abdomen densely pollinose, except for small nonpollinose dark spots around setae and in most a rather distinctive median dark vitta, which may be inconspicuous and only in posterior view indicated on the medio-dorsal region of T3; Tl +2 without median marginal setae; T3 with 4 strong marginal setae; genitalia (4 dissections): cerci and surstyli similar to those of ingerae n.sp. (Fig. 11); distiphallus very distinctive (Fig. 38). Female: Differs from the male as follows: face about 1.2x as long as frons; antenna about as long as face, segment 3 much narrower than maximal width of front femur; genitalia (3 dissections): St7 with short and weakly sclerotized apodeme (as in Fig. 8), only found in rossica of other European Siphona; St8 similar to that of rossica, but only with 5-7 setulae. Material examined: Finland: 6 d 4 Q, Karelia borealis. Sweden: 2 S?, Scania and Halland.

Distribution. A very rare and local species in North Europe. According to Mesnil (1965: 874) previously only known from a few specimens from Belgium, Germany and USSR (Type locality). Also recorded from Japan (Mesnil 1968: 163) and ? Mongolia (27 d 18 Y ? paludosa recorded by Herting 1968: 54, 1973b: 17).

Acknowledgements: My sincere thanks are extended to Dr. L. Lyneborg and to Dr. V. Michelsen, both of the Zoological Museum, Copenhagen, for advice and encouragement throughout the study and for construc- tive criticism of the manuscript. 1 am indebted to Dr. Brian H. Cogan and other colleagues in the British Museum (Nat. Hist.), London for commenting on the manuscript and for correcting my English, to Dr. B. Herting, Museum fur Naturkunde, Ludwigsburg, for valuable information and for loan of specimens, to Mr. O. Martin, Zoological Museum, Copenhagen who col-

lected Siphona in Finland, and to Dr. H. Andersson and Dr. R. Danielsson who supplied me with material from the collections of the Zoological Institute, Lund.

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Revision of European species of Siphona Meigen (Diptera: Tachinidae)

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