key: cord-0010283-w0lk3oya
authors: WHITE, IAN M.; KORNEYEV, VALERY A.
title: A revision of the western Palaearctic species of Urophora Robineau‐Desvoidy (Diptera: Tephritidae)
date: 2008-01-02
journal: Syst Entomol
DOI: 10.1111/j.1365-3113.1989.tb00289.x
sha: e064e42aa44b64fb41638dca48e2c2720111b44c
doc_id: 10283
cord_uid: w0lk3oya

Abstract A key is provided to twenty‐four western Palaearctic species of Urophora Robineau‐Desvoidy. The hosts of twenty‐three species which attack Asteraceae are listed, including those being used or investigated as possible weed biocontrol agents. The species are divided into four species groups and the differing host relationships and types of galls induced by these groups are discussed. U.lopholomae sp.n. and U.affinis ssp. calcitrapae ssp.n., associated with Centaurea (Lopholoma) spp. and C. (Calcitrapa) spp. respectively, are described. U. algerica (Hering) and U.sjumorum (Rohdendorf) are both treated as subspecies of U. quadrifasciata (Meigen). U.pontica is given full specific status and U.hispanica is removed from synonymy. The following new synonymies are made (junior synonyms in parentheses): U. angustifascia (Hering) (=Euribia phaeocera Hering); U. cardui (Linnaeus) (=U. reaumurii Robineau‐Desvoidy, lectotype designated); U. jaceana (Hering) (=E.conyzae Hering); U.maura (Frauenfeld) (=E. tecta Hering); U. mauritanica Macquart (=U. lejura Rondani, Trypeta macrura Loew); U.solstitialis (Linnaeus) (=E.sonderupi Hering, U. veruata Rondani); U.stylata (Fabricius) (=E.pia Hering, U. vulcaanica Rondani); U. terebrans (Loew) (=E. approximata Hering, T. eriolepidis Loew, E. manni Hendel). The possibility that U. quadrifasciata is a species complex is discussed; it is also suggested that U.affinis and U.jaceana represent the morphological extremes of a complex. The misuse of the name Musca stylata Fabricius in the genus Myopites Blot is noted.

been kept in the genus, and it is his restricted interpretation of the genus which is still applied today. The only named subgenus is Asimoneura which was described by Czerny (in Czerny & Strobl, 1909) for the Spanish species, U.stroblii Czerny. Between 1909 and 1963 the earlier name of Euribia was used by authors such as Hendel (1927) and Hering (1933 Hering ( , 1935 Hering ( , 1937 Hering ( , 1938a Hering ( , b, 1940 Hering ( , 1941 Hering ( ,1953 Hering ( ,1956 Hering ( ,1961 . However, in 1963 the International Commission for Zoological Nomenclature (opinion 678) ruled that the pamphlet of Meigen (1800) be suppressed and all species described as Euribia should therefore be placed in Urophora. The recent Catalogue of Palaearctic Diptera (Foote, 1984) listed sixtyfive species, thirty-three of which occur within the area covered by this paper. Since completion of the catalogue a further species has been described from this area by Freidberg (1982) ; Korneyev (1984 Korneyev ( , 1985 has added four species from the U.S.S.R., and single species have been described from Japan (Ito, 1983) and Korea (Kwon, 1985) . The present paper describes one additional species and a subspecies, and reduces the western Palaearctic list to twenty-five species as a result of new synonyms

The first key covering a substantial proportion of the genus was produced by Hendel(l927) and this was added to by Hering (1933 Hering ( ,1937 Hering ( ,1938a Hering ( , b, 1956 Hering ( ,1961 . The only modern work on the genus was compiled by Steyskal(l979) who presented separate keys for the Palaearctic and New World species of Urophora. However, Steyskal's work was not intended to be a revision and he discussed the need for a thorough revision of the genus. Steyskal (1979) drew attention to' the tentative nature of the section of his key that included many of the species of interest for the biocontrol of noxious weeds, sych as U.sirunaseva and U.solstitialis. An interim attempt to resolve this problem was made by White & Clement (1987) who discussed and keyed the species being used or tested as biocontrol agents for introduction into North America.

Traditionally, Urophora species have been identified primarily by characteristics of their wing patterns , together with femur, palp and antenna colour differences. Hering (1940) was the first author to utilize the shape of Fig. 6 , U.solstitiulis, dorsal view of hypandrium and aedeagal apodeme (stippled). Scale line=0.2 mm. aed. apod.. aedeagal apoderne: hypd., hypandrium.

the apex of the aculeus, or ovipositor piercer, as an aid to species diagnosis, but he seldom used the method in subsequent papers referring to the genus. Steyskal(l979) illustrated the aculeus tip shape for a few species, but not enough to enable the accurate identification of many of the critical species. In this paper, aculeus apex shape is used as a major identification character (Figs 30-64 ). Most species have very similar male terminalia. with neither the surstyli (Figs 1-5,7, 8, 10, 1 I ) , or the poorly sclerotized aedeagal glans yielding tenable diagnostic characters. However, a few species have well-sclerotized aedeagal glans and male terminalia characters can be used to divide the genus into four species groups; the critical characters are presented in the group diagnoses. Another male character that is sometimes species specific is the shape of the abdominal sternites . These vary from being very broad in the stem gall forming U.curdui (Fig. 21 ), to very narrow in U.angustifascia and both of the Echinops associated species . However, other species are very variable and have a sternite shape between those two extremes (Figs 17. 18) , although there may sometimes be a deep cleft in the most heavily sclerotized part of sternite 5, sometimes approaching the depth shown in Fig. 22 .

Most of the terminology used is that of White (1Y88) which was based on the system of McAlpine (1981) ; however, the term distiphallus has been corrected to aedeagal glans (Foote & Steyskal, 1987) . The terms primary and sccondary aculeus steps are also used; an aculeus with a single pair of preapical steps or shoulders is said to have primary steps only (e.g. Figs [43] [44] [45] ; an aculeus with two pairs of steps has both primary (proximal) and secondary (distal) steps (e.g. Figs 52-55). Following Steyskal (lY79) . the wing crossbands are named subbasal (covering bm-cu), discal (covering rm), preapical (covering dm-cu) and apical; see for examples with all four crossbands. Measurements were only taken from dissected females, and these are given in the species descriptions. with dimensions of exceptionally small or large individuals given in parentheses; malcs tend to smaller than females. The following abbreviations are used for measurements:

A L =Aculeus length (millimeters). W L P AL/WL= Aculeus length divided by wing =Wing length of female (millimetres) length.

A comprehensive study of larval and pupal stages is beyond the scope of this paper. However, pupal characters for most of the British species of Urophora have been tabulated by White (1988) . Published larval descriptions are catalogued by Hennig (1968) and Ferrar (1987) , and these authors also tabulate some larval characters.

The eggs of many tephritids can be dissected from museum-preserved adult females. Unfortunately this is rarely possible with Urophora species, suggesting that their eggs are generally less sclerotized than those of other Tephritidae. Eggs of U.stigma and U. hispanica were found in dry preserved specimens, but those of U.affinis and U.jaceana could only be extracted from specimens preserved in alcohol. It was found that the eggs of U.stigma are about 0.6 mm long, very broad and have a distinct apical opening (Fig. 24) which is presumed to be a micropyle (see Margaritis, 1985) . The eggs of the other species examined are much narrower and 0.6-0.8 mm long (Fig. 23 ). In the species groups proposed below, U.stigma is placed in a different group to the other three species whose eggs were examined, and this suggests that egg structure may differ between the groups.

The genus was first divided into species groups by Dirlbekova & Dirlbek (1980) who recognized three groups, namely 'stigma', 'cardui' and 'solstitialis', based on types of wing pattern. These authors also presented a key to the 'stigma' species group of Urophora which have hyaline rather than patterned wings. However, the present paper rejects this system of species groups in favour of one based on terminalia characters.

Each of the species groups defined in the present paper appears to be associated with a different tribe of the composite subfamily Asteroideae, as follows: group 1 includes all except three of the Palaearctic species and all those of known biology are associated with the Cardueae; group 2 is U.maura which is associated with h u l a species (Inuleae); group 3 includes U.stroblii whose host-plant is unknown and most, perhaps all, of the Afrotropical species; group 4 is U.stigma which is believed Western Palaearctic species of Urophora 333 to attack several species of Anthemideae; other undefined species groups include the New World species. These species groups are not formally named here because many of the eastern Palaearctic species have not been examined. Consequently, the old subgeneric limits are retained here, but they will probably be altered in a forthcoming study of the eastern Palaearctic species by the present authors.

Group 1 (subgenus Urophora in part): Species with known hosts are all associated with species of Cardueae (=Cynareae). Some species are known to spend the first larval instar in the egg and emerge as second instars; these are U.cardui (Peschken & Harris, 1975) , U.jaceana (Varley, 1937) and U.stylata (Redfern, 1968) . Species which are known to emerge as first instar larvae are U.afinis and U.quadrifasciata (P. Harris, pers. comm.) . Of these species, those that hatch as second instar larvae induce multilocular galls, while those that hatch as first instar larvae induce unilocular galls (P. Harris, pers. comm.) . Group 1 species of known biology induce a variety of gall types, usually formed of receptacle tissue (Zwolfer, 1983; Harris & Myers, 1984) , which engulfs the larval chamber that is situated in the achene or the receptacle (Freidberg, 1984) . These galls are lignified and may be multilocular, e.g. U.jaceana (Varley, 1947) and U.styZata (Harris & Wilkinson, 1984) or unilocular, e.g. U.afinis (P. Harris, pers. comm.) , U.jaculata and U.sirunaseva (White & Clement, 1987) ; often, unilocular galls occur multiply within a single flower head and they can be confused with the true multilocular galls. Conversely, U.quadrifasciata forms a papery, non-lignified, gall from the ovary wall (Harris &Myers, 1984) . The only species known to form a gall which is not within the capitulum of its host-plant, is U.cardui which forms a multilocular stem gall (Lalonde & Shorthouse, 1982 ) on a narrow range of Cirsium species. It is likely that galltype could be a useful classificatory character, but details of this kind are not yet available for most species. Members of this group are found throughout the Palaearctic region, with a single species in Taiwan, U. formosana (Shiraki) , and introduced species in North America and Australia.

Group 2 (subgenus Urophora in part): U.maura is the only known member of this group and it is associated with I d a species (Inuleae). Both the male and female terminalia of U.maura show a greater similarity to Myopites species than to other Urophora species; see illustrations of Myopites aculeus and aedeagal glans shapes given by Freidberg (1980) and White (1988) . In particular, the aedeagal glans of U.maura is a simple long tube with a sclerotized area near its base (Fig. 16) , very similar to known Myopites species; however, U. maura lacks the apical 'cup' shaped structure that is typical of Myopites species. Both Umaura and members of the closely related genus Myopites induce galls in the capitula of species of Inuleae and it is possible that U.maura is cladistically at least as closely related to Myopites species as it is to other Urophora species.

Group 3 (subgenus Asimoneura in part): U.stroblii is the only confirmed member of this species group, but its biology is unknown. This species has a very long labellum, a black scutellum and hyaline wings. The known Afrotropical species of Urophora also have these features and some, possibly all, of them may belong to this group. The only Afrotropical species dissected, U.petioZata Munro, was found to be similar to U,stroblii in terms o f the terminalia characters given in the group diagnosis, although the 'arms' of its aedeagal apodeme were broader. Many of the Afrotropical species are associated with Vernonia (Vernonieae) (A. Freidberg, pers. comm.) ; an exception is the Afrotropical U.pantomelaena Bezzi which develops in the capitulum of Helichrysum setosum (Inuleae) (Munro, 1926) and this may prove to belong to group 2. It is also likely that U.shirakii (Munro) from Taiwan belongs to this species group.

Group 4 (subgenus Asimoneura in part): U.stigma is the only known member of this species group and it forms a single non-lignified unilocular gall in a capitulum of an Achillea species (Anthemideae). Old host records suggest that it may also develop in the capitula of Anthemis and Chrysanthemum, which is likely because these genera also belong to the composite tribe Anthemideae.

Possible groups 5 onwards: an examination of the New World species of Urophora is beyond the scope of this paper. However, an examination of the key and descriptions produced by Steyskal (1979) suggests that none of the species native to the New World belong to the above species groups, and it is likely that all of the New World's native species belong to one or more as yet unrecognized species groups which a future reviser may decide to regard as separate genera. Little is known of their host-plants, but species of known biology appear to be associated with composite tribes such as Astereae and Heliantheae, neither of which are known host-plant groups for Old World Urophora species.

The following key uses external characters as far as possible. Host plants are used as key characters so that host-associated specimens of most species can be provisionally identified without the need for dissection. However, identifications made in this way should always be confirmed by dissection of female specimens. The major divisions in the key are based on external characters, 1 24 U 23 27 such as wing pattern, and the colour of the first flagellomere and femora. Specimens with aberrant wing patterning are fairly common and these will not run correctly in the following key. However, the alternative method of producing a key to this genus is to use aculeus tip shape characters for the major divisions in the key.

Such a key might be slightly more reliable, but it does necessitate dissection of all species, even if they are almost always easily distinguished by simple external characters. Confirmatory characters are given in square brackets and the descriptions should be consulted for other details such as measurements. Identification by aculeus tip shape is facilitated by the figures of the aculeus tips (Figs 30-64 Aculeus without preapical steps (Fig. 37) , or with indistinct steps (Fig. 36 

First flagellomere yellow or orange. Subbasal and discal crossbands usually separated by a yellow or hyaline area (similar to Fig. 66) ; if subbasal and discal crossbands joined (some U. aprica) then basal scutellar setae based in the black lateral areas of the scutellum, and aculeus apex with distinct preapical steps (Figs 48, 49 Hendel, 1927: 38 . Urophora (Urophora); Foote, 1984: 140. Subgeneric diagnosis. Lower facial margin not protruding; fold of proboscis and palpi not extending beyond oral margin; labellurn 1-2.5 times as long as first flagellomere; scutellum yellow centrally, with lateral black margins.

Urophora species group 1 Group diagnosis. Labellum usually about 1.5 times as long as first flagellomere, except for species associated with Echinops (U.pontica and U.syriaca) in which it is about 2.5 times as long as first flagellomere. Scutellum yellow, black at sides. Aedeagal glans reduced to a narrow membranous sac, with no heavily sclerotized areas . 'Arms' of aedeagal apodeme broadly joined to hypandrium (Figs 1, 4-6). Spermathecae not sclerotized (it is not usually possible to find them).

Comments. It is possible that a careful study of male terminalia characters might allow some further division of this group. For example, the only stem gall former is also the only species with remarkably broad maler sternites (Fig. 21) . Similarly, some of the species with pointed aculei (Figs 40, 41) , also have very narrow male sternites and the aedeagal glans of U.pontica and U.syriaca (Fig. 14) differ from other species (typically as in Figs 12, 13). U.pontica and U.syriaca are the only species associated with Echinops, and they also differ from other species by having very broad palpi (Figs 27,28), a longer labellum, and by the black basal area of their wings. These data suggest that the two Echinops associated species could be regarded as a distinct species group, and that would concord with a recent cladistic study of the tribes of Asteraceae (Bremer, 1987) which places Echinops in a separate tribe (Echinopsideae) from the Cardueae.

Trypeta afJinis Frauenfeld, 1857: 541 . Urophora afjinis (Frauenfeld) ; Zwolfer, 1965: 138; Steyskal, 1979: 14; Foote, 1984: 140. Comments. The only available specimen which is almost certainly a syntype of U.afJinis is a male, so it cannot be confirmed that it is the species which attacks Acrolophus in Central Europe. However, a female which is likely to be a syntype, was dissected and the identity of this species is assumed to have been confirmed by examination of that specimen. Urophora afJinis is here divided into two subspecies, which not only differ in their morphology, but also in their distributions and host associations. The two subspecies may be separated using the following key:

1 Wing length (9) usually less than 3.4 mm. Aculeus length usually less than 1.9 mm, without deep primary steps (Figs area covered by this paper. However, in the Ukrainian S.S.R. it has been reared from Centaurea (Phaloiepis) sterilis, a species which is known to hybridize with some Acrolophus species (J. Dostal, in Moore et al., 1976) , for example C. diffusa and C. maculosa, both of which are known hosts of U.ajjinis ssp. afinis. This subspecies emerges from its egg as a first instar larva and the larvae induce unilocular galls within the host capitulum (P. Harris, pers, comm.) .

Comments. lJ.afinis and U.jaceana have very similar aculeus tip shapes and the possibility that these two species are very closely related is discussed later in this paper. This subspecies has been established in North America as a biocontrol agent for Centaurea diffusa and Centaurea sp. near maculosa (Harris & Myers, 1984; Piper, 198.5 Biology. This subspecies is only known to attack species of Centaurea subgenus Calcitrapa and it is known from several countries between Turkey and Iran.

Comment. It is possible that the two 'subspecies' of U.affinis are distinct species, and the evidence for this is given in the discussion

Holotype, 9 , ex Centaurea iberica, ISRAEL: Mount Hermon, 1600 rn, 2.vii.1986 (emerged by 17.ix.1986) (Freidberg) (BMNH); not dissected, but aculeus tip exposed.

Paratypes. Ex C. calcitrapa, TURKEY: 3 9 , 1.5 km N Balikesir, 20.vii.1984 N Balikesir, 20.vii. (emerged by 24.v.1985 

Euribia angustifascia Hering, 1956: 83 . Euribia phaeocera Hering, 1961 : 319 syn.n. Urophora angustifascia (Hering) ; Steyskal, 1979: 9; Foote, 1984: 141. Redescription. Head: first flagellomere black; palpi orange in specimens from Turkey and Israel, but darkened to grey or black apically in specimens from Iran and Afghanistan; palpi shape similar to U.cardui; gena about 0.3 times eye height; labellum about 1.5 times length of first flagellomere. Thorax: scutum with a dense tomentum which obscures the underlying cuticle; postpronotal lobe largely yellow; basal scuttellar seta based on the central yellow area; marginal black area of scutellum very narrow; femora black, except for extreme apex which is orange. Wing: base hyaline or yellow; subbasal crossband extending between veins C and A , ; subbasal and discal crossband joined between veins C and R,; discal and preapical crossbands separate; hyaline area between preapical and discal crossbands 2.5 times as broad along vein R4+5 as breadth of preapical crossband on vein R4+5; preapical and apical crossbands joined between C and R2+3 or R4+>. Terminalia: aculeus apex, BioIogy. U. angustifascia has been reared from Cousinia hermonis in northern Israel (Freidberg, 1974b) . U.angustifascia is also known from Turkey, Iran and Afghanistan and Cousinia is a very diverse genus in these areas (Wagenitz. in Davis, 1975; Wagenitz, 1980) , so it is likely that Cousinia species are the normal hosts of this species. Furthermore, U.angustijmcia belongs to a Middle-Eastern species group all of which have a pointed aculeus and black femora, and the only other member of this group with a recorded host is U.melanocera (Hering) , which is also associated with a Cousinia species (Korneyev, 1983) .

Comments. U.angustifascia and U.phaeocera were separated by Hering (1961) and Steyskal (1979) largely on the basis of palpi colour differences. These two nominal species have identical aculeus shapes and they are very similar in all other respects, so they are here assumed to be conspecific. Should a future reviser find good evidence that they are in fact distinct species, it should be noted that both Hering (1961) and Steyskal (1979) erroneously reversed the palpi colours of these nominal species in their keys.

Holotype, 0 , E.angustifascia, IRAN: Baluchistan, Iranshar; not located. Paratype, E. angustifascia, IRAN: 9 (not dissected), data as holotype (BMNH). Holotype, d (not dissected), E.phaeocera, ISRAEL: (TAU). Other material: 1 6 , 1 9 , Turkey, Adana, Karatas (BMNH); I ? , Afghanistan, Badakschan (BMNH).

Tephritis aprica Fallen, 1820: 7. ? Urophora brunicornis Robineau-Desyoidy, 1830: 772. Urophora centaureae (Fabricius, 1794) ; Robineau-Desvoidy, 1830: 772 [misidentification]. Urophora scutellata Rondani, 1970: 15,21 . Urophora aprica (Fallen) yellow; basal scutellar seta based within the broad marginal black area, except in some Turkish specimens; femora black, at least in basal two-thirds orange apically, except in Turkish specimens which have extensive orange areas on at least the fore-and mid-femora. Wing: base yellow; subbasal crossband extending between veins C and A1; subbasal and discal crossbands separated by a narrow yellow area, but sometimes joined between veins C and R4+5; discal and preapical crossbands separated; hyaline area between preapical and discal crossbands 1- Comments. The name U.aprica has been misapplied to small specimens of U. terebrans (previously known as U.approximata) and to U.lopholomae. Although it is a Fallen species, Hering (1938b) actually described U. approximata as a new species for what he thought was the 'aprica' of Fallen and then gave a key to separate it from the 'aprica' of Loew. The lectotype is mould covered and in a delicate state. Consequently, it was not dissected, but its external features are sufficiently distinct as to leave little doubt about the identity of this species. U. brunicornis was described within Robineau-Desvoidy's (1830) description of U. centaureae suggesting that it is likely to be a synonym of U.aprica.

Lectotype, 9 (designated by Persson, 1958;  not dissected), T.aprica, SWEDEN: (NRS). (Linnaeus) ; Steyskal, 1979: 12; Foote, 1984: 141. Redescription. Head: first flagellomere orange; palpi orange; palpi shape, Fig. 29 ; g e m about 0.25-0.5 times eye height; labellum about 1-1.5 times length of first flagellomere. Thorax: scutum with a dense tomentum which obscures the underlying cuticle; postpronotal lobe largely yellow or orange; basal scutellar seta on or near margin of central yellow and marginal black area; femora usually black in basal two-thirds, sometimes black and orange striped, rarely dark orange. Wing: Fig. 68 ; base yellow; subbasal crossband extending between veins C and hind margin of wing; subbasal and discal crossbands joined between veins C and R4+5; discal and preapical crossbands joined between middle of cell dm and hind margin of wing; hyaline area between preapical and discal crossbands, measured along vein R4+5, approximately equal in breadth to preapical crossband on vein R4+5; preapical and apical crossbands joined between C and R4+5. Terminalia: aculeus apex, Fig. 43 ; 8 terminalia, Figs 1,2,12. 6 sternites 4 and 5 , Fig.   21 . WLP=4.9-5.4 mm; AL=1.7-1.9 mm; AL/ WL=0.304).40.

Biology. U.cardui is only known to attack Cirsium arvense and C. creticum. The larvae emerge from the egg in the second instar and they induce a multilocular stem gall. No other Urophora species is known to induce a stem gall and it has the shortest aculeus length to wing length ratio of any European species. However, the eastern Palaearctic species U.misakiana, U. bicoloricornis and U. hoenei each have a similarly short aculeus, which is almost identical to U.cardui in its apical shape. Unfortunately, the biology of these species is unknown, but they may also prove to be stem gall formers.

Comments. The lectotype of U.cardui (designated by White, 1987) , is only known through an illustration (Reaumur, 1738) , and the same illustration is referred to in the original description of U. reaumurii; the lectotype of U.cardui is here designated as the lectotype of U.reaumurii, so these nominal species are objective synonyms. Although this lectotype, and the syntypes of other synonyms have all been lost, Reaumur (1738) illustrates both the morphology and biology of U.cardui in great detail and there is no doubt that it refers to the species which forms stem galls on Cirsium arvense. U.cardui has been established at a few sites in Canada and the U.S.A. for the biocontrol of Cirsium arvense (Peschken er al., 1982; Piper, 1985; Story, 198%) .

Cirsium arvense: 3 Q , 16, England, three localities (BMNH); 1 9 , France, Caen (BMNH); 1 $2, Germany, Berlin (BMNH); 1 9 , Hungary, Velence (BMNH). C.creticum: 1d,2P, Greece, Kavalla (BMNH). No host data: 1 9 , Yugoslavia, Macedonia, Lake Ochrid, (BMNH). (Fig. 44 ) Urophora congrua Loew, 1862: 74; Steyskal, 1979: 9; Foote, 1984: 141. Redescription. Head: first flagellomere orange; palpi orange; palpi shape similar to U.cardui; gena about 0 . 3 4 . 4 times eye height; labellum about 1-1.5 times length of first flagellomere. Thorax: scutum with a dense tomentum which obscures the underlying cuticle; postpronotal lobe largely yellow; basal scutellar seta on or near margin of central yellow and marginal black area; femora orange, often extensively marked with black, sometimes with fore-and hind-femora almost entirely black in basal half. Wing: base hyaline; subbasal crossband extending between veins C and A2,; subbasal and discal crossbands joined between veins C and R4+5; discal and preapical crossbands joined between C and R4+5; hyaline area between preapical and discal crossbands 0.75-1.5 times as broad along vein R4+5 as breadth of preapical crossband on vein R4+5; preapical and apical crossbands joined between veins C and R4+s. Terminalia: aculeus apex, Fig. 44 . W L P = 4 . 4 5 . 4 mm; AL=2.2-2.8 mm; AL/WL=0.45-0.55.

Biology. This species attacks the capitula of Cirsium erisithales (Zwolfer, 1965) .

Comments. The aculeus tip shape of U.congrua (Fig. 44) is very similar to that of U.stylata (Fig. 4 3 , suggesting that these two species associated with Cirsium species may be closely related, or similarly adapted for oviposition in the capitula of that genus.

Presumed syntypes, AUSTRIA: I d (not dissected), 1 P (aculeus tip broken) (no label data) (Egger) (ZMHB); the original description implies that many more syntypes from Bavaria and Austria were examined by Loew, but these have not been located. Other material, Cirsium erisithales: 129, 2 8 , Austria, three localities (SMNS, ZMUC).

Urophora cuspidata (Meigen) (Fig. SO Fig. 50 . WLE =4.@-6.0 mm; AL=(2.8-) 3.1-4.2 mm; AL/WL=0.554.75.

Biology. Attacks the capitula of Centaurea subgenus Lopholoma species and the larvae induce multilocular galls within the capitulum.

Comments. Although the only possible syntype could not be positively identified because its ovipositor is broken, the size and wing pattern of this specimen is typical of the species associated with Centaurea subgenus Lopholoma. apically; palpi shape similar to U.cardui; gena about 0.3-0.4 times eye height; labellum about 1.5 times length of first flagellomere. Thorax: scutum with a dense tomentum which obscures the underlying cuticle; postpronotal lobe largely yellow; basal scutellar seta on or near margin of central yellow and marginal black area, except in alpine specimens (associated with Centaurea alpestris) which have a broader black margin to the scutellum so that the basal seta is based within the black area; femora orange, usually striped black. Wing: base yellow; subbasal crossband extending between veins C and A,; subbasal and discal crossband separated by a yellow area; discal crossband often reduced to spots in British specimens, sometimes reduced to a single mark along the r-m crossvein; discal and preapical crossbands separate; hyaline area between preapical and discal crossbands 1.5-2 times as broad along vein R4+5 as breadth of preapical crossband on vein R4+5; preapical and apical crossbands usually joined between C and (BMNH); 2 9 , 16, England, two southern localities (BMNH); 109, 1 6 , France, four southern localities (not reared) (BMNH); 1 0 , Germany, Naumberg on Saale (BMNH). No host data: 2 9 , Norway, Akershus, Berum, Ostoya (ZMUB).

Urophora species nr cuspidata (Meigen) (Fig. 51) Diagnosis. Similar to U.cuspidata, except in the following: femora orange; suhbasal crossband extending between veins C and A2, sometimes almost reaching hind margin of wing; hyaline area between preapical and discal crossbands 1-2 times as broad along vein R4+s as breadth of preapical crossband on vein R4+5; preapical and apical crossbands joined between C and R2+3. Terminalia: aculeus apex, Fig. 51 , WL0=4.7-4.9 mm; AL=3.5 mm; AL/WL= 0 . 7 O . 7 5 .

Biology. Two females of this species have been reared from the capitula of Centaurea (Cyanus) depressa.

Comment. The deep apical notch and primary steps of this species (Fig. 51) indicate that it is probably a very close relative of U.cuspidatu (Fig. SO) , but the section beyond the primary steps is very elongate and it lacks the distinct sccondary steps of U.cuspidata. A single male of Terellia colon (Meigen) , or a very closely related species, was also reared from the same collection of C. depressa capitula. T. colon, like U . cuspidata, is normally only associated with Centaurea subgenus Lopholoma; this implies that C. depressa is probably very similar to Lopholoma species in terms of the characteristics that are of importance to these tephritids. This Urophora species is not formally described here because the aculeus tip shape could only be studied on one specimen and the possibility that this is simply an aberrant U.cuspidata cannot be discounted.

Centaurea depressa: 2P ( 1 with aculeus tip broken off). Turkey, Konya, Beysehir, junction of roads to Konya, lsparta and Seydesehir, 17.vii. 1984 (emerged by 15.v. 1985 ) (Rosenrhul) (USNM) . (Fig. 46) Urophora hermonis Freidberg, 1974a: 49. Redescription. Head: first flagellomere orange; palpi yellow, darkening to orange apically; palpi shape similar to U.cardui; gena about 0.4 times eye height; labellum about 1.5 times length of first flagellomere. Thorax: scutum with a dense tomentum which obscures the underlying cuticle; postpronotal lobe largely yellow; area of scutum ajoining postpronotal lobe and notopleuron yellow; basal scutellar seta based within the central yellow area; femora black in basal half, then mostly orange. Wing: lacking crossbands; orange infuscate, usually more deeply orange anterior to vein &+s than rest of wing. Terminalia: aculeus apex, Fig. 46 . WL?=4.6-5.0 mm; AL=3.4-3.8 mm; AL/ WL=0.70-0.80.

Biology. Induces galls in the capitula of Cousinia hermonis (Freidberg, 1974b) . Urophora hispanica Strobl (Figs 23, 36, 37) Urophora (afinis ?ssp.) hispanica Strobl, 1906: 358. Urophora uffinis (Frauenfeld, 1857) ; Zwolfer, 1965: 141 [misidentification] . Urophora sp. nr aSfinis (Frauenfeld, 1857) ; Zwolfer, 1965: 146 [misidentification] . Urophora algira Macquart, 1843; auctt. (in part) [misidentifications].

Redescription. Egg, Fig. 23 . Head: first flagellomere orange; palpi yellow, darkening to orange apically; palpi shape similar to U.cardui; gena about 0.3 times eye height; labellum about 1.5 times length of first flagellomere. Thorax: scutum with a dense tomentum which obscures the underlying cuticle; postpronotal lobe largely yellow; basal scutellar seta based in yellow area, or at margin of central yellow and marginal black U.algira. The latter species was described from a male which was 2 lines (4.2 mm) long. As males of U.hispanica are only about 3 mm long it is unlikely that the name U.algira applies to this species. area; femora yellow or orange. Wing: base hyaline; subbasal crossband extending between veins R1 and A,, or absent; discal and preapical crossbands separate; hyaline area between preapical and discal crossbands 1.5-2 times as broad along vein R4+5 as breadth of preapical crossband on vein %+5; preapical and apical crossbands separate. Terminalia: aculeus apex, Figs 36, 37. WL?=2.7-3.5 mm; AL=1.4-1.9 mm ; WL/AL= 0.45-0.60.

Biology. Urophora hispanica attacks several distantly related species of Centaureinae, namely Mantisalca salmantica and starthistles belonging to the Centaurea subgenera Seridia, Solstitiaria and possibly Calcitrapa.

Comments. The syntypes of U. hispanica could not be examined, but it was described as being very similar to U.afjinis, and the type locality is in coastal north-eastern Spain. The only species known from the western mediterranean coastal area, which resembles U.afinis, is this species which attacks Mantisalca and starthistles. Consequently, the name U. hispanica is assumed to apply to this species. This species has previously been misidentified as U.afJinis and

Centaurea aspera: 9 9 , 2 8 , France, nine southern localities (BMNH, LTUB, SMNS); I ? , Spain, Granada, Lobres (not reared) Hering, 1935: 169 . Euribia solstitialis (Linnaeus, 1758); auctt. pre 1935 (in part) [misidentification]. Urophora solstitialis (Linnaeus, 1758); auctt. pre 1935 (in part) [misidentification]. Urophora jaceana (Hering, 1935) ; Steyskal, 1979: 20; Foote, 1984: 142. Redescription. Head: first flagellomere orange; palpi yellow, darkening to orange apically; palpi shape similar to U.cardui; gena about 0.4-0.6 times eye height; labellum about 1-1.5 times length of first flagellomere. Thorax: scutum with a dense tomentum which obscures the underlying cuticle; postpronotal lobe largely yellow; basal scutellar seta on or near margin of central yellow and marginal black area; femora orange, often with black areas or streaks. Wing: Fig. 66 ; base hyaline to yellow; subbasal crossbands extending between veins C and A l , or C and AZ; subbasal and discal crossbands separated by a yellow area; discal and preapical crossbands separate; hyaline area between preapical and discal crossbands 1-3 times as broad along vein R4+s as breadth of preapical crossband on vein R4+,-; preapical and apical crossbands usually joined between C and R2+3, or C and R4+s, but sometimes separate. Terminalia: aculeus apex, Figs 60, 61. W L 9 = 3.2-4.6 mm; AL=2.0-2.6 mm; A L / W L = 0 . 5 0 . 7 0 .

Biology. U.jaceana attacks the capitula of knapweeds belonging to Centaurea subgenus Jacea in which it induces a lignified multilocular gall of receptacle tissue (Varley, 1947) , and the larvae emerge during the second instar (Varley, 1937) . In southern France a population of Urophora which were morphologically identical to U . jaceana were swept off C . (Acrolophus) spinabadia at a site where no subgenus Jacea could be found. Acrolophus species are the normal hosts of U.afinis ajj'inis, which has an identical aculeus tip shape to U.jaceana, suggesting that these may be closely related species. The possibility that U.ajj'inis ujj'inis and U.jaceana are simply the morphological extremes of a complex of closely related and very similar species, is discussed later in this paper.

Comments. U.conyzae is the oldest available name for the knapweed gall fly. However, the case 26801. name U.jaceana is well known because of the lifetable study of this species carried out by Varley (1947) , which is still quoted to students of population ecology. Consequently, an application has been made to the International Commission for Zoological Nomenclature to have the name U. jaceana given precedence over U. conyzae by any authors who regard them as synonyms (Case 2680). U.juceana is also known from North America where it has been accidentally established since before 1923 (Shewell, 1961) .

Syntypes, E.jaceana, ex Centaurea jacea, GERMANY / POLAND border: 1 9 , River Oder at Crossen, 3.vii.1934 (reared) (Hering) (BMNH); 7 6 , 7 9 (none dissected), same locali ty , 10.vi. 193421 .vi .1935 (Hering) (BMNH) .

FRANCE: 1 9 , 1 6 (not dissected), Paris, Lardy, 18.vii.1932 (not reared) (Hering?) (BMNH). Other material, Centaureajacea: 5 9 , France, Pyrenees-Oriental, Font-Romeu (BMNH); 2 6 , 3 9 , Germany, same data as syntypes (BMNH); 1 9 , Hungary, Pilis, Dobogoko (not reared) (BMNH). C.nigra: 9 9 , 188, England, seven localities (BMNH). C.phrygia, 1 9 , Austria, Eisenstadt (SMNS). C.spinabadiu: 8 9 , France, Pyrenees Oriental, Collioure, Tour Madeloc (not reared, but no other likely host at site, 45 not dissected) (BMNH) [Plant determined by G. Wagenitz] . No host data: 2 9 , Norway, Akershus, Berum, Ostoya (ZMUB); 1 ? , Romania, Transylvania (BMNH); 2 9 , Scotland, Badenoch & Strathspey, Grantown (BMNH). (Figs 60,6!, 66) Urophora jaculata Rondani, 1870: 14, 18 . Urophora sirunaseva (Hering, 1938b) ; auctt.

[misidentification].

Head: first flagellomere orange; palpi yellow, darkening to orange apically; palpi shape similar to U.cardui; gena about 0.4 times eye height; labellum about 1.5 times length of first flagellomere. Thorax: scutum with a dense tomentum which obscures the underlying cuticle; postpronotal lobe largely yellow; basal scuteller seta on or near margin of central yellow and marginal black area; femora . black area; femora black, except at apex. Wing: base hyaline to yellow; subbasal crossband extending between veins C and A,; subbasal and discal crossbands separated by a yellow area; discal and preapical crossbands separate; hyaline area between preapical and discal crossbands about 0.75 times as broad along vein R4+5 as breadth of preapical crossband on vein R4+5; preapical and apical crossbands joined between C and R4+5. Terminalia: aculeus apex, Fig. 47; 8 terminalia, Figs 3 , 4 , 13. W L 9 =3.4-3.8 mm;

AL=1.6-1.7 mm; AL/WL=0.40-0.45.

Diagnosis. This species is easily distinguished. from other Urophora species, including the superficially similar U.aprica, by its characteristic aculeus tip shape (Fig. 47) .

Biology. The type host of U.lopholomae is Centaurea sadleriana, but a single male which is probably this species was swept from a plant identified as C x a b i o s a . These two plants are very closely related species of subgenus Lophnloma, and it is possible that the plant identified as the common and widespread

Comment. This species has previously been confused with U.aprica.

orange. Wing: base hyaline to yellow; subbasal crossband extending between veins C and AZ; subbasal and discal crossbands separated by a yellow area; discal and preapical crossbands separated; hyaline area between preapical and discal crossbands about 0.7.5 times as broad along vein R4+5 as breadth of preapical crossband on vein R4+s; preapical and apical crossbands joined between C and R4+5. Terminalia: aculeus apex, Figs 38, 39. W L Q =3.2-3.5 mm; AL=1.5-2.0 mm; AL/WL=0.504.60.

Biology. U.jaculata attacks Centaurea solstitialis in Italy and in mainland Greece, except in the north-east where this plant is attacked by U.sirunaseva. U.jaculata induces lignified unilocular galls in the capitulum of C.solstitialis (White & Clement, 1987) and it has also been swept from another member of Centaurea subgenus Solstitiaria.

Comment. This species used to be confused with U.sirunaseva and all references to that species in pre-1987 weed biocontrol literature actually refer to U.jaculata (White & Clement, 1987) .

Lectotype, Q (designated by White & Clement, 1987), ITALY: no label data, but distribution given in original description as Parma countryside and more southern provinces (MZF). Other material, Centaurea ? napifolia: 2 9 , Italy, Catanzaro (not reared) (SMNS). C.solstitialis: 4 9 , Greece three mainland localities (BMNH); 8 9 , 2 8 , Italy, five localities from Rome to Sicily (BMNH). Urophora lopholomae sp.n. (Figs 3 , 4 , 13 (Fig. 42) ?Urophora algira Macquart, 1843: 378 . Urophora mauritanica Macquart, 1851: 259 . Trypeta macrura Loew, 1855 : 40 syn.n. Urophora lejura Rondani, 1870: 14, 19 syn.n. ?Urophora sejuncta Becker, 1907: 390 . Urophora sp. nr solstitialis (Linnaeus, 1758) ; Zwolfer, 1965: 146 [misidentification] .

Urophora macrura (Loew, 1855) ; Foote, 1984: Urophora sibynata Rondani, 1870 ; auctt.

[misidentification].

Head: first flagellomere orange; palpi yellow, darkening to orange apically; palpi shape similar to U.cardui; gena about 0.3-0.4 times eye height; labellum about 1-1.5 times length of first flagellomere. Thorax: scutum with a dense tomentum which obscures the underlying cuticle; postpronotal lobe largely yellow; basal scutellar seta on or near margin of central yellow and marginal black area; femora orange, with a black stripe on at least the forefemur. Wing: base hyaline to yellow; subbasal crossband extending between veins C and A,; subbasal and discal crossbands separated by a yellow area; discal and preapical crossbands separate; hyaline area between preapical and discal crossbands usually 1.5-2 times as broad along vein R4+5 as breadth of preapical crossband on vein R4+5, but up to 4 times in some areas of North Africa; preapical and apical crossbands separate, or joined, often as far as vein &+5; preapical and apical crossbands reduced to spots in some areas of North Africa. Terminalia: aculeus apex, Fig. 42 . WLP (2.943.5-5.6 mm ; AL= (2.5-)3. >5.4(-6.0) mm ; Biology. In Carthamus arborescens, U.mauritanica induces lignified multilocular galls. It also attacks the capitula of other Carthamus species, but it has not been recorded from the, commercially grown safflower, Ctinctorius L.

Comments. The oldest name which may apply to this species is U.algira. However, its type was not found amongst Macquart material in either MNHNP or UMO, and it is probably in MHNL, where most of the surviving Macquart material is badly mould damaged. The name U.algira has previously been applied to U.hispanica, but the described size of the male of U.algira is larger than U . hispanica. The only large species which is known from Algeria is the Urophora species associated with Carthamus. The oldest name which definitely applies to this species is U. mauritanica , and U. macrura and U. Iejura are synonyms. U.sejuncta from Tunisia is probably a synonym of U. mauritanica because it is the only large Urophora species known from North Africa. AL/WL=0.75-1.00(-1 .lo).

Syntype, U.mauritanica, ALGERIA: 1 9 , no label data (UMO). Syntypes, T.macrura, GREECE: 19 (not dissected but aculeus tip exposed), 16 (not dissected) (Kiesenwetter) (ZMHB) . Syntypes, U.lejura, ITALY: 1 0 , 4 8 (not dissected), Apennine mountains near Parma (as 'Apenninis montuosis ditionis parmensis') no label data (MZF). Other material, Carthanus arborescens: 6 9 , 2 8 , Spain, single localities in Granada and Malaga provinces (BMNH). Clanatus: 3 9 , France, three southern localities (LTUB, BMNH). Ctenuis: 2 9 , Israel, two localities (BMNH). No host data: 1 P , Cyprus, Zakaki (BMNH); 1 0 , Italy, Sardinia, Gairo (BMNH); 1 0 , Yugoslavia, Macedonia, Prespa Geul, Otesevo (BMNH); 2 9 , Libya, Tripolitania, Horns (BMNH); 1 0 , Malta (BMNH); 1 9 , Morocco, Ito (BMNH); 2 9 , Algeria, Dazinville (BMNH).

Syntype, U.algira (6 only, mentioned in description), ALGERIA: (presumed to be in MHNL); not available for examination. Syntypes, U.sejuncta, TUNISIA: near Tunis, v. 1906 (Becker); not located. (Fig. 30) Urophora neuenschwander Freidberg, 1982: 56. Redescription. Head: first flagellomere black, sometimes dark orange on inner side; palpi yellow, darkening to orange or grey apically; palpi shape similar to U.cardui; gena about 0.3 times eye height; labellum about 1-1.5 times length of first flagellomere. Thorax: scutum with a fine tomentum which does not obscure the underlying cuticle; postpronotal lobe yellow laterally, but black on the medial side of the postpronotal seta; basal scutellar seta on or near margin of central yellow and marginal black area; femora black, except for the apex which is orange. Wing: base yellow; subbasal crossband extending between vein C and the hind margin; subbasal and discal crossbands joined from C to R4+5; discal and preapical crossbands separate; hyaline area between preapical and discal crossbands less than half as broad along vein crossband extending between veins C and A,; subbasal and discal crossbands joined between C and R4+5; discal and preapical crossbands separate; hyaline area between preapical and discal crossbands 0.75-1 times as broad along vein R4+5 as breadth of preapical crossband on vein R4+5; preapical and apical crossbands joined, usually to R4+5. Terminalia: aculeus apex, Fig Comments. Previous authors have called this species U.dzieduszyckii, but that name refers to a much larger species whose female was described as having a wing length of 4.4 mm. Specimens from France were regarded as the nominal subspecies of U.dzieduszyckii by Hering (1937), who described subspecies pontica from near Volgograd, because it lacked the abnormally long fore coxae of his nominal subspecies. Hering (1937) described the nominal subspecies as having the fore coxae as long as the fore femur. In fact, Hering's own specimens (BMNH) have the fore coxae 0.74.8 (36) and 0.8 (1 9 ) times as long as thefore femur, and the specimens labelled as 'paratype ssp. pontica' in the BMNH have the fore coxae 0.6 ( I ? , 16) times as long as the fore femur (this is typical of all other Urophora species); an additional specimen from Turkey has a ratio of 0.8 (19). These data suggest that there may be a tendency towards abnormally long fore coxae in Mediterranean and western European individuals of this species, but the evidence is tenuous and there is no good reason to give any of these poorly known populations subspecific status.

R4+5 as breadth of preapical crossband on vein R4+5; preapical and apical crossbands joined from C to %+5. Terminalia: aculeus apex, Fig.  30 , WL?=3.5-3.6 mm; AL=2.2 mm; AL/ WL=0.60-0.65.

Biology. U.neuenschwanderi attacks the capitula of Ptilostemon species, belonging to the section of the genus that are shrubs, rather than herbs. It is only known from Crete and the type host is Ptilosternon gnaphaloides (Freidberg, 1982) , but there is also a single rearing record from Staehelina arborea (Neuenschwander & Freidberg, 1983) ; Xarborea forms part of the same chasmophytic plant community as the shrubby Ptilostemon species and this record probably represents an oviposition error by U. neuenschwanderi. U. neuenschwanderi is only known from Crete and its competition with other species attacking the same host is discussed by Neuenschwander (1984) .

Holotype, 0 (not dissected), ex Ptilostemon gnaphaloides, GREECE: Crete, Chania Province, Aerinos, 18.vi. 1980 (emerged 16.v. 1981 ) (Neuenschwander) (TAU). Paratypes, ex P.gnaphaloides, GREECE: 3 d (one labelled allotype, none dissected), 2 9 , Crete (TAU). Other material, P.chamaepeuce: 1 6 , 2 9 , Greece, Crete, Nio Horio (not reared,,but no other likely host at site) (BMNH). Urophora pontica Hering 5tat.n. (Figs 5, 14, 20, 28, 41) Euribia dzieduszyckiipontica Hering, 1937: 244 . ffrophora dzieduszyckii Frauenfeld, 1867;  auctt.

[misidentification].

Head: gena sometimes black below the eye; first flagellomere dark orange to grey; palpi black; palpi, Fig. 28 ; gena about 0.4-0.5 times eye height; labellum about 2-2.5 times length of first flagellomere. Thorax: scutum with a fine tomentum which does not obscure the underlying cuticle; postpronotal lobe yellow laterally, and black on the medial side of the postpronotal seta; basal scutellar seta on or near margin of central yellow and marginal black area; femora dark orange to black, except for apex which is black. Wing: base black; subbasal

Syntypes, U.S.S.R.: 1 9 (not dissected), 16 (not dissected), Russian SFSR, near Volgograd, Sarepta (Becker 36581) (8 BMNH, 9 ZMHB); specimens labelled as types, but not listed in original description, 2 9 (one in BMNH dissected), Sarepta (Becker 33610 Urophora quadrifasciata (Meigen, 1826) ; Stey-772. skal, 1979: 15; Foote, 1984: 143. Comments. Urophora quadrifasciata is a very variable species and two forms of aculeus shape can be distinguished (Figs 31,32, & 33-35) . One of these forms can be further divided by differences in aculeus length. The resulting three forms each attack non-overlapping ranges of host-plants and have some geographic separation. There is some rather circumstantial evidence that U. quadrifasciata is a species complex, and this is presented in the discussion. With the evidence available at present, it would be unwise to attempt to distinguish true species within this possible complex. However, the three morphologically distinct forms may be regarded as subspecies and these are separated by the following key: Aculeus margins swollen before apex; each margin is not parallel to the lumen wall (Figs 33-35) .

[AL= 1.9-3.0 mm. Aculeus apex at least 8 Fm wide. Aculeus usually less than 2.0 mm long (AL=1.3-2. I mm); sample means less than 2.0 mm (Fig. 31) .

Associated with knapweeds (mostly Centaurea subgenera Acrolophus and Jacea) and Centaurea solstitidis. ............. quadrifasciata quadrifasciata Aculeus usually more than 2.0 mm long (1.8-3.0 mm); sample means greater than 2.0 mm (Fig. 32) .

Associated with Centaurea subgenus Calcitrapa, namely C.calcitrapa in Turkey, and C.iberica in Israel and Pakistan; specimens without host data have been found in Armenian S.S.R., Cyprus, Iran and Turkmen S.S.R.). .. quahifasciala sjurnorurn Urophora quadri fasciata quadrifasciata (Meigen) 5tat.n. (Figs 31,67) Redescription: Head: first flagellomere yellow to black; on Centaurea arenaria grey in 62% of specimens examined, yellow in 25% and black in 6% (n=8); on Cdiffusa grey in 83%, black in 11%, yellow in6% (n=19); on C.maculosa grey in 73%, black in 22%, yellow in 5% (n=56); on C.nigra grey in 53%, black in 47% (n=34); on C.solstitialis grey in 82%, black in 18% (n=9); palpi yellow, darkening to orange apically; palpi shape similar to U.cardui; gena about 0.254.3 times eye height; labellum about 1.5 times length of first flagellomere. Thorax: scutum with a fine tomentum which does not obscure the underlying cuticle; basal scutellar-seta on or near margin of central yellow and marginal black area; femora black, except at extreme apex. Wing: Fig. 67 ; base yellow; subbasal crossband extending between veins C and A, or between C and AZ; subbasal and discal crossbands joined between C and R4+5; discal and preapical crossbands separate; hyaline area between preapical and discal crossbands usually about 0.75-1.5 times as broad along vein f&+5 as breadth of preapical crossband on vein R4+5; preapical and apical crossbands joined between C and R4+5. Terminalia: aculeus apex, Fig. 31 . Biology. U. quadrifasciata quadrifasciata attacks the capitula of Centaurea species belonging to subgenera Acrolophus, Cyanus, Jacea, Phalolepis and Solstitiaria. In Acrolophus species which are adventive in Canada, this subspecies of U.quadrifasciata induces a thin nonlignified gall, formed from the wall of an ovary (Harris & Myers, 1984) , and the larvae emerge in the first instar (P. Harris, pers. cbmm.). Sobhian & Zwolfer (1985) describe the gall of U. quadrifasciata in Centaurea solstitialis as 'a gall in the ovary and later [the larva] feeds within a transformed achene the seed coat of which is inflated'. This reference to an inflated seed coat suggests that the galls examined by Harris & Myers (1984) and Sobhian & Zwolfer (1985) may differ in structure, but this has not been confirmed.

Comments. U. quadrifasciata quadrifasciata is established in North America on Centaurea sp. near maculosa and Cdiffusa (Harris & Myers, 1984; Piper, 1985; Story, 1985a) .

Syntypes. T. quadrifasciata, GERMANY: 1 8 (not dissected), 1 9 (not dissected), ('Muhlheim am Rhein') no label data (MNHNP). Other material, Centaurea alba: 2 9 , Italy, Rome (BMNH). Carenaria: 1 6 , 9 9 (Figs 33-35) Euribia algerica Hering, 1941: 52 . Urophora algerica (Hering, 1941) ; Steyskal, 1979: 15; Foote, 1984: 140. Redescription. First flagellomere grey (n= 15 off Cuspera), except in 12% of specimens off Cnicaeensis, black (n=9); aculeus apex, Figs 33-35; WL9=2.4-3.8 mrn; AL=1.9-3.0 mm; apex breadth at least 8 pm (see key); ALI WL=0.60-0.80; other details as nominal subspecies.

Biology. The known hosts of U . quadrifasciata algerica are starthistles belonging to the Centaurea subgenera Seridia and Solstitiaria.

Comments. Hering (1941) described Urophora algerica on the basis of two male specimens, so it is not possible to confirm the identity of this nominal species. However, its wing pattern is typical of U. quadrifasciata, although Hering (1941) compared it to U.congrua. The only available female specimen from Algeria which has a 'quadrifasciata' type wing is clearly related to the southern Italian Urophora quadrifasciata population which attacks Centaurea nicaeensis and this implies the identity of U. algerica.

Holotype, 6 (not dissected), ALGERIA: between Blida and Medea, 7.viii.1884 (Quedenfeldt) (ZMHB); one wing of the holotype, mounted on a slide (BMNH); specimen labelled as a type and mentioned in original description, but not listed as a type, ITALY: Calabria (Erber) (ZMHB). Other material, Centaurea' aspera: 2 8 , 1 8 9 , Spain, Granada Prov., Lobres (not reared) (BMNH). Cnicaeensis: 7 9 , Italy, two Puglia localities (BMNH). N o host data: 1Q , Algeria (BMNH).

(Rohdendorf) stat.n. (Fig. 32) Euribia sjumorum Rohdendorf, 1937: 144. ? Euribia armeniaca Hering, 1937: 245 . Urophora sjumorum (Rohdendorf, 1973) ; Steyskal, 1979: 15; Foote, 1984: 144. Redescription. First flagellomere yellow in 60%, grey in 40% of specimens (n= 17); aculeus apex, Fig. 32 ; WL9 =2.6-4.1 mm; AL=2.0-3.0 mrn; AL/WL=0.654.85; other details as nominal subspecies.

Biology. U.quadri&asciata sjumorum has been reared from the capitula of plant species belonging to Centaurea subgenus Calcitrapa in Turkey, Israel and Pakistan; specimens were also examined from Cyprus and Iran, and the types of U.sjumorurn and its probable synonym were collected in Turkman S.S.R. and Armenian S.S.R.

Comments. It is clear from the original description of U.sjumorum that it is a form of U.quadrifasciatn with a long ovipositor; its type locality indicates that it is almost certainly the form associated with Centaurea subgenus Calcitrapa. Hering (1937) describes U. armeniaca as being similar to U.hoenei, which is a Chinese species that is closely related to U.cardui, but he says the ovipositor is longer than the abdomen. The available slide mounted wing labelled as 'armeniaca' in the Hering collection (BMNH) has the typical pattern of U.quadrifasciata; its size (WL=4.1 mm), long ovipositor and type locality, indicated that it is also most likely to be this subspecies of U. quadrifasciata.

Presumed syntype (could be part of holotype or paratype), E.armeniaca, U.S.S.R.: wing on slide, Armenian S.S.R. (BMNH). Other material, Centaurea calcitrapa: 4 9 , Turkey, two localities (USNM). Chyalotepis: 18, 4 9 , Israel, Benot Ya'aqov Bridge (BMNH). Ciberica: 18, 219, Israel, Mount Hermon (BMNH). No host data: 5 9 , Cyprus, Cherkes (BMNH); 1 9 , Pakistan, Mingosa (BMNH).

Holotype, Q , E.sjumorum, U.S.S.R.: Turkmen S.S.R.; not located. Holotype, ? , E.armeniaca, U.S.S.R.: Armenian S.S.R. (in coll, 'Hermann, Erlangen'); not located. Paratype, E.armeniaca, U.S.S.R.: 1 9 , data as holotype (presumed to be with holotype).

Eurihia satunini Zaitzev, 1945: 379 . Urophora satunini (Zaitzev, 1945); Foote, 1984: 144.

Comments. The syntypes of this nominal species were not located and the original description, written in Georgian, was not available in any library accessible to us. Consequently this species is not included in the key. It was described from north-east Turkey, close to the border with Armenian S.S.R.. and it is hoped that it will be possible to include Umtunini in a later paper which will cover the Soviet species (Korneyev & White, in prep.) .

Chaldyr; not located.

Urophora sirunaseva (Hering) (Fig. 52) Euribia sirunaseva Hering, 1938b: 397 . Urophora ulgira Macquart, 1843; Steyskal, 1979: 17 [misidentification] . Urophora sirunaseva (Hering, 1938b) ; Steyskal, 1979: 18; Foote, 1984: 144. Redescription. Head: First flagellomere orange; palpi yellow, darkening to orange apically; palpi shape similar to U.cardui; gena about 0.3 times eye height; labellum about 1.5 times length of first flagellomere. Thorax: scutum with a tomentum which barely obscures the underlying cuticle; postpronotal lobe largely yellow; basal scutellar seta on or near margin of central yellow and marginal black area; femora orange. Wing: base hyaline to yellow; subbasal crossband extending between veins C and A,; subbasal and discal crossbands separated by a yellow area; discal and preapical crossbands separate; hyaline area between preapical and discal crossbands usually about 2 times as broad along vein R4+5 as breadth of preapical crossband on vein &+s, except in Crete and Israel where the separation is only about equal to the breadth of preapical crossband on vein &+5; preapical and apical crossbands separate, except in Crete and Israel where they are joined almost to R4+5. Terminalia: aculeus apex, Fig. 52 . WLP =(2.6-)2.9-3.7 mm; AL=-i.4-2.l--mm; ALiWL=0.50-0.65.

Solstitiaria. In north-eastern Greece, Turkey, Moldavian S.S.R., Ukrainian S.S.R. and northern Israel it is associated with C.solstitiafis, and in Crete it attacks the closely related Cidaea. In both of these hosts the larvae induce lignified unilocular galls, which often occur multiply.

Comment. Records of this species from Italy and other areas of Greece refer to U.juculata; this misidentification caused confusion to the weed biocontrol programme against C.solstitialis in North America (White & Clement, 1987) .

Syntypes, ex Centaurea solstitialis, U.S.S.R.: 18, 1 9 (neither dissected), Moldavian S.S.R., Bendery (as 'Tighina') (BMNH); the following specimens labelled 'paratypes', but not listed in the original description: 3 8 (not dissected), 49

(not dissected), 1 9 , same data as syntypes (Figs 6,17, Rondani, 1870: 14, 18 . Euribia sonderupi Hering, 1940 : 1 syn.n. Urophora sp. nr approximatu (Hering, 1938b) ; Zwolfer, 1965: 95 [misidentification] . Urophora solstitialis (Linnaeus, 1758) ; Steyskal, 1979: 14; Foote, 1984: 144. Redescription. Head: first flagellomere orange; palpi yellow, darkening to orange apically; palpi shape similar to U.cardui; gena about 0.4-0.5 times eye height; labellum about 1-1.5 times length of first flagellomere. Thorax: scutum with a dense tomentum which obscures the underlying cuticle; postpronotal lobe largely yellow; basal scutellar seta on or near margin of central yellow and marginal black area, except in some individuals from mountain areas which 1830: 770.

have the setae based within the marginal black area; femora orange with dark stripes, except in some individuals from mountain areas which have extensive black marks on the femora and in some cases almost completely black femora. Wing: base hyaline to yellow; subbasal crossband extending between veins C and A,; subbasal and discal crossbands separated by a yellow area; discal and preapical crossbands separated; hyaline area between preapical and discal crossbands about 2 times as broad along vein R4+5 as breadth of preapical crossband on vein R4+5; preapical and apical crossbands separate, or sometimes joined, but rarely beyond R,.

Terminalia: aculeus apex, Fig. 53 ; aedeagal apodeme, Fig. 6 . WL9 =(2.7-)3.&4.8 mm; AL= (1.7-)2.3-3.9 mm; AL/WL=0.604.90; the specimens from Denmark (reared from Cirsium) are at the lower end of the size range.

Biology. U.solstitialis normally attacks the capitula of some Carduus species, namely C.acanthoides, C.crispus and C. nutans, in which the larvae induce lignified multilocular galls (White & Clement, 1987) . In the mountain areas of the Jura, the Alps and southern Germany, a form with dark femora and broad black margins to the scutellum has been found associated with C.defloratus; there is no evidence that this is not conspecific with other Carduus associated populations and the dark coloration may be the effect of cold winter diapause; such an effect has been experimentally induced in some butterflies such as Pieris brassicae (Gardiner, 1979) . Furthermore, alpine populations of U.cuspidata have similar broad black margins to the scutellum. U.sonderupi is here regarded as a synonym of U.solstitialis, although it was apparently reared from Cirsium vulgare, rather than from a species of Carduus; this unusual host association is detailed in the discussion.

Comments. No female syntype of U.solstitialis could be located, but Linnaeus described this species as being Carduus associated and its identity is assumed from that information (White, 1987) . Similarly, the types of U.dauci, U.leucacanthi and U. femoralis were unavailable for study, and the types of U.hastatus and U.pugionata were too frail to be dissected; for nomenclatorial stability these nominal species, whose types have the typical wing pattern and size of U.solstitialis, were all assumed to be synonyms of U.solstitialis. Both U. veruata and U.sibynata have the typical aculeus tip shape and wing pattern of the species which attacks Carduus and they are also assumed to be that species. U.sonderupi also has this aculeus tip shape and wing pattern, but it was apparently reared from Cirsium vulgare. 'paratype' (from Maribo)(BMNH), are not syntypic because they do not match the data in the original publication; 16, 3 9 from Norre Alslev are dated 22.vi.1941 which is 8 months after the description was published; specimens erroneously labelled as types, Lolland, Maribo, 29.iii-1 .iv. 1935 and 12.vii-22.viii. 1940 ('inflor .-Galle') (Sonderup) (BMNH); 1 0 , Maribo, 24.viii.1940 (? reared) (Sonderup) (BMNH). Other material, Carduus acanthoides: 3 P , Austria, Sollenau (not reared) (BMNH); 2 ? , Germany, Upper Palatinate, near Regensburg (BMNH).

C.crispus, 1 9 , France, Alsace, Hirtzfelden (BMNH). C.dejZoratus: 1 9 , Austria, Tirol.

Otztal, Zwieselstein (SMNS); 7 9 , Switzerland, Jura and three localities in Valais (BMNH, SMNS, ZMUC). C.nigrescens: 1 6, 1 0 , France, Herault, Salvetat (BMNH). C.nurans: 1 0 .

England, Surrey, Banstead Downs (BMNH); 4 9 , 1 6 , France, three localities (BMNH); 1 9.

Germany, Neuenberg (BNMH); 1 9 , Turkey, Erzurum (USNM); 1 9 , U.S.S.R., Ukrainian S.S.R. (ZIL). Cirsium helenioides: 2 9 , Switzerland, Valais, Gletch (SMNS). No host data: Form with dark femora, known to be associated with Carduus dejloratus, 1 9 , Germany, Alterburg (BMNH).

Other type data lieved lost.

Syntypes, T. leucacanthi, 'Ingolstadt': be-Urophora spoliata (Haliday) (Fig. 54) Tephritis spoliata Haliday, 1838: 186 . Urophora spoliata (Haliday, 1838) ; Steyskal, 1979: 12; Foote, 1984, 144. Redescription. Head: first flagellomere orange; palpi orange; palpi shape similar to U. cardui; gena about 0.3 times eye height; labellum about 1-1.5 times length of first flagellomere. Thorax: scutum with a dense tomentum which obscures the underlyingcu-tide; postprontal lobe largely yellow; basal scutellar seta usually based within the marginal black area; each femur mostly black, but often not on all faces, even in the basal half. Wing: without crossbands; cell sc yellow; rest of wing hyaline. Terminalia: aculeus apex, Fig. 54 . W L ? =3.2 mm; AL=1.7-2.0 mm; AL/WL=0.5&0.65.

Biology. U.spoliata attacks the capitula of Serratula tinctoria in a small area of southern England; there are no confirmed records from any other areas.

Probable syntype, ENGLAND: 1 9 (not dissected), Hants, Isle of Wight (no label data) (NMID). Other material, Serratula tinctoria: 1 6 , England, Hants, Leckford (BMNH) ; several other reared specimens from' Leckford and Cranmore, Isle of Wight, seen (BMNH), but not dissected; 3 9 , 18, not reared, dissected.

Urophora styhta (Fabricius) (Figs 45,65) Musca stylata Fabricius, 1775 : 785. Trupanea cirsii Schrank, 1808 : 140. Musca jacobeae Panzer, 1805 : 22. Urophora venabulata Rondani, 1870 : 14, 17. Urophora vulcanica Rondani, 1870 Euribia pia Hering, 1938a : 244 syn.n. Urophora stylata (Fabricius, 1775) ; Steyskal, syn.n. 1979: 13; Foote, 1984: 144 .

Redescription. Head: first flagellomere orange; palpi yellow, darkening to orange apically; palpi shape similar to U.cardui; gena about 0.25-0.4 times eye height; labellum about 1.5 times length of first flagellomere. Thorax: scutum with a dense tomentum which obscures the underlying cuticle; postpronotal lobe largely yellow; basal scutellar seta on or near margin of central yellow and marginal black area; femora orange. Wing: Fig. 65 ; base hyaline; subbasal crossband absent, or rarely extending between veins R, and A l ; discal and preapical crossbands separate; hyaline area between preapical and discal crossbands 1.5-2 times as broad along vein R4+5 as breadth of preapical crossband on vein R4+5; preapical and apical crossbands usually Biology. This species probably occurs throughout the Palaearctic region, and it has been found in Pakistan. It attacks the capitula of Cirsium species in most of Europe, and it has been swept from a Cirsium species in Japan. However, in the Mediterranean area it has been found associated with Carduus and Galactites species (Neuenschwander & Freidberg, 1983) . In Cirsium vufgare its larvae induce a multilocular gall (Harris & Wilkinson, 1984) and its larvae emerge during the second instar (Redfern, 1968) .

Comments. The type of U.stylata lacks an abdomen, so even if it is a female the identity cannot be confirmed. However, the type locality is England, and the wing pattern of the available syntype is typical of the species which attacks Cirsium vulgare in England. The identity of the available syntypes of U. venabulata was confirmed by examination of the aculeus apex. U.pia was described by Hering (1938a) from a single male which was collected at the same locality, and by the same collector, as typical specimens of U.stylata, and U.pia is probably only an aberrant U.stylata that differs from typical specimens by having a partly formed subbasal crossband. Similarly, U. vulcanica could only be identified by its wing pattern and no syntypes of U.cirsii or U.jacobeae could be located; these species were regarded as synonyms of U.stylata to maintain nomenclatorial stability. The eastern Palaearctic species U.suchalinensis (Shiraki), U.japonica (Shiraki) and U.carnpes-tris Ito are very similar to U.stylata, and their status will be discussed in a later paper (Korneyev & White, in prep.) . U.styfata has been established in North America for the biocontrol of Cirsium arvense (Harris & Wilkinson, 1984; Piper, 1985) . The name Musca stylata Fabricius 1775, has also been used for a species of the related genus Myopites Blot, normally known as 'M.styfatus (Frabricius, 1794)'; Fabricius (1794) gives Musca stytata as a redescription from his work of 1775. Only Urophora stylata fits the original description of Musca stylata which was described from England, an area from which the species known as 'Myopites styfatus' is not known to occur (White, 1988) ; this Myopites species should therefore be known by its synonym Myopites limbardae Schiner, 1864 .

Syntype Musca stylata, ENGLAND: 1 (no abdomen) ('in Angliae nemoribus') no label data (ZMUC). Syntypes, U. venabulata, ITALY: 36 (not dissected), 5 9 (not dissected, but 2 with exposed aculeus), 2 (no abdomens), ('in tota Italia') no label data ( 

? Urophora dzieduszyckii Frauenfeld, 1867: Euribia syriaca Hendel, 1927: 49 . Euribia erichischmidti Hering, 1953: 2. Urophora syriaca (Frauenfeld, 1867) ; Steyskal, 498. 1979: 9; Foote, 1984: 145. Redescription. Head: first flagellomere dark orange to grey; palpi black; palpi shape, Fig. 27 ; gena about 0.34.6 times eye height; labellum about 1.5-2.5 times length of first flagellomere. Thorax: scutum with a fine tomentum which does not obscure the underlying cuticle; postpronotal lobe yellow laterally, black on the medial side of the postpronotal seta; basal scutellar seta on or near margin of central yellow and marginal black area; femora black, except at extreme apex. Wing: base black; subbasal crossband extending between veins C and A1 or AZ; subbasal and discal crossbands joined between C and R4+5; discal and preapical crossbands separate; hyaline area between preapical and discal crossbands up to 2 times as broad below vein R4+5 as breadth of preapical crossband on vein R4+5, but narrower than preapical crossband above R4+5; preapical and apical crossbands joined from C to R4+5. Terminalia: aculeus apex similar to Fig. 41 ;d sternites 4 and 5, Fig. 22 . WL? =3.8-4.3 mm; AL=2.4-2.7 mm ; AL/WL=0.55-0.75.

Biology. U.syriaca is only known from the Lebanon, plus nearby areas of Syria and Israel. In northern Israel it has been reared from an Echinops species (Kugler & Freidberg, 1975) .

Comments. The oldest available name that may apply to this species is U.dzieduszyckii, but this nominal species has no type locality and the female from which it was described could not be located. Previous authors, such as Hendel (1927) and Foote (1984) , give the type locality of U. dzieduszyckii as being Poland. This is an error caused by the fact that Frauenfeld (1867) was sent the specimen by M. Nowicki of Krakow, Poland. Frauenfeld also acknowledges W. Dzieduszycki, who was presumably the collector, and notes that Dzieduszycki was a supporter of a museum in Lemberg, now in Latvian S.S.R. Hendel(l927) examined the type of U. dzieduszyckii, and even presents a photograph of its wing. Frauenfeld described the species as being larger that U.eriolepidis (a synonym of U.tere-brans), and as having a wing length of 4.4 mm. It is just possible that U.dzieduszyckii is an aberrant U.cardui found in Latvian S.S.R., but the wing photograph presented by Hendel (1927) and its large size, suggest that U.dzieduszyckii is more likely to be the Echinops associated species found in the Middle East. There is little doubt that the holotypes of U.syriaca and U. erichischmidti also represent this Echinops associated species, which is here referred to as U.syriaca as that is the oldest confirmed name. Shammay (not reared) (BMNH); 1 0 , 2 d , without any host data, from Israel, also dissected.

Holotype, 9 , U. dzieduszyckii, LOCALITY UNKNOWN: not found; however Hendel (1927) figures its wing.

Urophora terebrans (Loew) (Fig. 55) Trypeta terebrans Loew, 1850: 53 . Trypeta eriolepidis Loew, 1856 : 52 syn.n. Euribia manni Hendel, 1927 : 45 syn.n. Euribia approximata Hering, 1938b : 398 syn.n. Urophora aprica (Fallen, 1820) ; auctt. partim Urophora terebrans (Loew, 1850); Steyskal, [misidentification]. 1979: 17; Foote, 1984: 145. Redescription. Head: first flagellomere orange; palpi yellow, darkening to orange apically; palpi shape similar to U.cardui; gena about 0.2-0.5 times eye height; Labellum about 1-1 .5 times length of first flagellomere. Thorax: scutum with a dense tomentum which obscures the underlying cuticle; postpronotal lobe largely yellow; basal scutellar seta on or near margin of central yellow and marginal black area; femora black, at least in basal two-thirds, sometimes with dark orange stripes. Wing: base hyaline to yellow; subbasal crossband extending between veins C and A , or A2; subbasal and discal Hardy, 1968 ), E.manni, AUSTRIA: no label data (NHMV). Paralectotype, E.munni, AUSTRIA: 1 9 , no label data (NHMV).

GERMANY: 1 6 , 1 9 , 'Beuron (Alb.) ', iii.1937 (Ziegenhagen) (BMNH). Other material, Comments. Previous authors, for example Steyskal (1979) , separated U.terebrans and U.eriolepidis on the basis of the relative breadth of the hyaline area between the discal and preapical wing crossbands; however, this appears to be geographic variation in a single species. Specimens from Germany, including the syntypes of U. approximata, are sma!ler than U.terebrans from other areas, but they have the same aculeus tip shape and they attack the same host as U.terebrans in the Pyrenees; these differences are probably also explained by geographic variation. Hendel (1927) described U. manni because he said it had 3-4 pairs of frontal setae, rather than the usual 2 pairs. In fact the type specimens each differ in the number of frontals on each side of their heads, and the paralectotype was found to have the typical aculeus tip shape of U.terebrans. Consequently, it is assumed that the types of U.manni are simply aberrant specimens of U. terebrans.

WL=0.55-1.00. Group diagnosis. Labellum about 1-1.5 times as long as first flagellomere. Scutellum yellow, black at sides. Female T6 shiny centrally, tomentose laterally. Spermathecae well sclerotized and ridged (Fig. 25 ). Aedeagal apodeme with narrow 'arms' (Figs 8, 9) . Aedeagal glans reduced to a long membranous tube with a distinct basal sclerite (Fig. 16) :

Urophora maura (Frauenfeld) 16, 25, 62) Trypeta maura Frauenfeld, 1857: 550 . Euribia tecta Hering, 1940: 2. syn.n. Urophora maura (Frauenfeld, 1857) ; Steyskal, 1979: 11; Foote, 1984: 143. Redescription. Head: first flagellomere orange; palpi yellow, darkening to orange apically; palpi shape similar to U. cardui, but slightly narrower; gena about 0.2-0.3 times eye height; labellum about 1-1.5 times length of first flagellomere. Thorax: scutum with a dense tomentum which obscures the underlying cuticle; postpronotal lobe often largely yellow, but sometimes mostly black with a small yellow spot just posterior to postpronotal seta; basal scutellar seta based within the central yellow area of the scutellum; femora orange. Wing: without crossbands; entirely hyaline except for yellow cell sc. Terminalia: aculeus, Fig. 62 and Richter (1970) lists Lsalicina as a host, but there is no indication of the original source of this record. Although no reared specimens were examined. there is no reason to doubt the association with Inula. The records of U.maura associated with Acroptilon repens (L.) DC. (Tyurebaev. 1981; Watson & Harris, 1984) probably refer to either U.xanthippe (Munro) or U. impicta (Hering) .

Comment. The holotype of U.tecta was dissected and its aculeus shape was found to be typical of U.maura. No tenable differences were found between these nominal species and they arc therefore regarded as synonyms. ; Hendel, 1927: 38. Urophora (Asimoneura) ; Foote, 1984: 145. Subgeneric diagnosis. Lower facial margin protruding; fold of proboscis and palpi extending beyond oral margin; labellum about 3 times as long as first flagellomere.

Group diagnosis. Labellum about 3 times as long as first flagellomere. Scutellum black. Spermathecae not sclerotized (it is not possible to find them). Aedeagal glans reduced to a narrow membranous sac, with no heavily sclerotized area. 'Arms' of aedeagal apodeme narrowly joined to hypandrium (similar to Fig. 9 ).

Urophora stroblii (Czerny) (Fig. 63) Asimoneura stroblii Czerny, in Czerny & Strobl, 1909: 253. Euribia (Asimoneura) stroblii (Ezerny, 1909) ; Hendel, 1927: 48 . Urophora (Asimoneura) stroblii (Czerny, 1909) ; Foote, 1984: 146. Redescription. Head: first flagellomere black; palpi yellow; labellum about 3 times length of first flagellomere. Thorax: postpronotal lobe black; scutellum black; femora and tibia black; tarsi yellow. Wing: without crossbands; cell sc yellow; rest of wing hyaline. Terminalia: aculeus, Fig. 63 . W L 8 = 2 . 0 mm; W L 9 = 2.4 mm; AL=1.08 mm; AL/WL=0.45.

Biology. U.stroblii is the only Urophora species included in this paper for which there is no available biological data.

Synt-ypes, SPAIN: 1 8 , 1 P , Gerona Province, Monistrol, 14-22.v. 1907 (data from original publication, n o label data) (Strobl) (NHMV),

Group diagnosis. Labellum about 3 times as long as first flagellomere. Scutellum yellow, black at sides. Female T6 tomentose. Spermathecae well sclerotized and covered in warty tubercles (Fig. 26 ). Aedeagal apodeme with narrow 'arms' (Fig. 11; similar to Fig. 9 ). Aedeagal glans largely comprised of a wellsclerotized basal section with a narrow membranous extension (Fig. 15 ).

during Nelson's bombardment of Copenhagen.

Urophora stigma (Loew) (Figs 10, 11, 15, 24, 26, 64) ? Muscaplacida Muller, 1764: 85 . Trypeta stigma Loew, 1840: 156 . Trypeta unimaculata Roser, 1840 : 60. Euribia (Asimoneura) stigma (Loew, 1840) ; Urophora (Asimoneura) stigma (Loew, 1840) ; Hendell927: 47. Foote, 1927: 145. Redescription. Egg, Fig. 24 (Schlechtendel, 1891) , but none of these records have been confirmed. In Achillea nobilis the larvae induce a nonlignified unilocular gall in the capitulum, which appears to be formed from the ovary wall in a similar manner to the gall of U. quadrifasciata in some Centaurea subgenus Acrolophus species.

Presumed syntypes, T.stigma, POLAND: 1 d (not dissected), Silesia (Scholtz) (ZMHB); LOCALITY UNKNOWN: 1 d (not dissected), labelled '626' (ZMHB). Syntypes, T. unimaculata, GERMANY: 1 (no abdomen) (SMNS); possible syntypes or author determined specimens: l ? , I d (neither dissected) (SMNS).

Other material, Achillea millefolium: 1 d , Germany, Sud-Kyffhauser (BMNH). A.nobilis: 1 d (not dissected), 1 gall, probably Austria (NHMV). No host data, eggs, 2 9 , 16, France, three southern localities (BMNH).

Many of the Urophora species described in this paper are very variable in characters such as size and wing pattern. Previous authors, such as Hendel(1927) and Hering(1937 Hering( ,1938a Hering( , b, 1940 Hering( , 1961 , accepted some mere varietal forms as distinct species. For example, U.terebrans and U.approximata were distinguished by the relative lengths of their ovipositors, despite the fact that both of these nominal species attack the same host and there is no evidence to suggest that the longer ovipositor of U.terebrans is anything other than an allometric artefact of its larger body size. Loew (1856) , and all subsequent authors, distinguished U. terebrans, from the Pyrenees, and U.eriolepidis, from Austria, by the relative separation of their discal and preapical crossbands, although there is no other distinct difference between these two nominal species. Similarly, Hendel (1927) separated U.manni from other Urophora species because he said it has three, rather than two pairs of frontal setae. In fact the lectotype of U.rnanni has three frontal setae on one side of its head, and two on the other, and it is clearly an aberrant individual of U. terebrans. These earlier authors were unaware of allometry, they did not have large enough collections with which to detect geographic variation in wing patterns, and they often failed to detect aberrant chaetotaxy. They also lacked the benefit of our present knowledge of host-plant relationships, which was largely founded upon the work of Zwolfer (19650, and they were unaware of the taxonomic value of the aculeus tip shape. In this paper U. eriolepidis, U.manni and U.approximata are all placed in synonymy with U.terebrans because there is no good evidence to suggest that they are distinct species, although it would be unwise to discount such a possibility entirely. A similar approach was taken in the interpretation of all the species covered by this paper. Consequently, the nomenclatural confusion caused by such illfounded nominal species as U.rnanni should have been overcome. However, some of the socalled 'species' described in this paper may in fact be groups of very closely related species that cannot be easily distinguished morphologically. The possible species complexes will now be discussed.

In central Europe U.afJinis subspecies afjinis and U. jaceana are only known to attack species of the Centaurea subgenera Acrolophus and Jacea, respectively. These two Urophora species are superficially quite different in appearance, as they differ in both size and wing pattern, but they share an almost identical aculeus tip shape (Figs 5 M 1 ) which suggests that they may be closely related species. In central Europe they are never found on each other's range of hosts and there is little doubt that they are distinct species in that area. However, individuals that are morphologically identical to U. jaceana have been found on a species of Acrolophus on the Mediterranean coast of France. Furthermore, flies which are the size of U.jaceana, but with the wing pattern of U.affinis, are found attacking species of the subgenus Centaurea (Calcitrapa) in Turkey and the Middle East, and these populations are named here as U.ufjinis subspecies calcitrapae. A few specimens reared from Acrolophus species in Turkey were examined, and found to be typical of subspecies afjinis. The discovery of both subspecies in an area of sympatry, suggests that the two 'subspecies' may in fact be distinct species. However, the only Turkish specimens of subspecies afjinis were not from sites where subspecies calcitrapae was found, so there is no evidence that the two subspecies are distinct when they occur together at a single site. Consequently, the two forms of U.affinis should be treated as subspecies until more conclusive evidence is obtained to support their separation. Similarly, the flies from southern France, which were found on Centaurea (Acrolophus) spinabadia, are unlikely to be genetically typical U.jaceana, as that species has never been found associated with such closely related plant species as C. ( A . ) maculosa, although C.maculosa has been well surveyed by several weed biocontrol specialists. These data suggest that central European U.afJinis subspecies affinzs and U.jaceana may simply be the morphological extremes of a series of at least four groups of populations, some or all of which may be reproductively isolated and therefore distinct species.

Urophora quadrifasciata, as delimited in this paper, has the longest host list of any species of Urophora, although all of its confirmed hosts belong to a single genus, namely Centaurea. R. D. Wild (unpublished CIE report, 1987) showed that populations associated with different hosts and geographic areas are morphometrically distinct. In this paper, three groups of populations are regarded as subspecies, each of which occurs within a section of the total distribution, and has a host list which does not overlap with the host lists of the other two subspecies. At the extreme ends of the variation in the aculeus tip shape of U. quadrifasciuta are Central European individuals of the nominal subspecies (Fig. 31) , and the single available Algerian female of U. quadrifasciata algerica (Fig. 33 ).

Between these two extremes, every conceivable intermediate form of aculeus shape has been found. However, U.quadrifasciata algerica could be further subdivided according to geographical origin or host (Figs 33-35) , and there is some tentative biological evidence that some populations of the nominal subspecies cannot attack some of the hosts listed for that subspecies. U. quadrifasciata quadrifasciata is widespread on adventive species of Acrolophus in North America, but it rarely attacks C. (Jacea) nigra, although that is a heavily attackedplant in Europe. Similarly, it has spread into the North American range of C.solstitialis, but it has not yet been recorded as attacking that plant, although that is a host of the nominal subspecies in Europe. Without biological evidence, such as the results of host choice tests, combined with the discovery of morphometric differences between sympatric populations, there is no good reason to regard any of the populations here referred to as U. quadrifasciata as anything other than forms of a single polytypic species. However, appropriate study may show that this socalled 'species' is actually a complex of several species, some or all of which only attack a very narrow range of closely related Centaurea species.

Urophora stylata attacks more than one hostplant genus, but it shows no obvious variation which correlates with the host differences. No detailed morphometric analysis has been carried out, but there is no indication that U.stylata populations associated with Cirsium species in central Europe are morphologically different from Mediterranean populations associated with Carduus and Galactites.

Urophora solstitialis attacks Carduus species found on Onopordon, Cynara and Carlina, lack any differences other than size, and there is no evidence that they are not conspecific.

The main aim of this study was to revise the species, concepts and nomenclature of western Palaearctic Urophora species so that a new key could be produced and a meaningful host-plant list compiled. In any genus of monophagous or oligophagous insects it is impossible to be certain about the true limits of species that show hostrelated variation, unless very expensive and detailed studies are undertaken of each of the potential species complexes. This revision has stopped short of actually attempting detailed studies of each possible species complex. As a compromise, the available evidence suggesting that some of the 'species' described here may actually be species complexes has been presented, and these data should be taken into account when using the host list (see appendix) as a reference source for selecting potential weed biocontrol agents. throughout most of its range, but from the Danish islands of Lolland and Falster there are some reared series from Cirsium vulgare. This plant has been well surveyed in many areas of Europe by Zwolfer (1965) , and by several subsequent weed biocontrol specialists, but no further finds of U.solstitialis have been made on it. However, there is a single record of specimens identical to normal U. solstitialis reared from Cirsium helenioides in Switzerland (leg. Zwolfer). The Danish specimens are smaller than typical Curduus associated specimens of U.solstitialis and they were described as a separate species called U.sonderupi by Hering (1940) . It is possible that U.sonderupi is a distinct species, but this is unlikely as it has never been found again on Cirsium vulgare. It is also possible that the host-plant was a misidentified Carduus species, possibly C.acanthoides or C.crispus, but that does not explain why the specimens were unusually small. It is also possible that Cirsium vulgare is the normal host of U.solstitialis in Scandinavia, as that area has not been well surveyed. No obvious conclusion can be drawn from these possibilities and U.sonderupi is here regarded as a synonym of U.solstitialis because there is no good evidence that it is a distinct species.

U. hispanica and U.terebrans are very variable species, both of which appear to have an ecologically determined host range. U. hisppnica attacks species of Centaureinae which have a spinose bract appendage, and which current botanical classifications separate into two genera, namely Mantisalca (previously Centaurea (Microlonchus)), and the Centaurea subgenera Calcitrapa, Seridia and Solstitiaria. All of the species attacked by I/. hispanica are component parts of plant communities which grow adjacent to each other in coastal western Mediterranean areas. This suggests that the flies associated with these plants are probably a single species which has a preference for particular habitats, as well as plant type. Similarly, U.terebrans attacks thistles with large capitula, regardless of whether they are Carlina, Cirsium, Cynara or Onopordon species. One of the largest forms, and the smallest form of U.terebrans, both attack Cirsium eriophorum; the large form is found in the Pyrenees and the small form in Germany (formerly known as U. approximata). In all respects other than size these two populations, plus Urophora specimens that have been

Texas A & M University), for their assistant in the collection of Urophora. Thanks are also due to professor H. Zwolfer (LTUB) for his helpful discussions of Urophora biology and for the loan of many valuable specimens; to the Godman Fund (BMNH) for kindly helping towards the cost of I.M.W.'s collecting in Spain and Washington State, U.S.A.; and A . C. Pont (BMNH) for reading a draft of the manuscript. The following curators kindly arranged loans of material in their charge: A. C. Pont (BMNH)

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Special thanks are due to the Ministry of Forests, British Columbia, for funding field trips for I.M.W. to collect tephritids in Europe, lsrael and Canada, that were of relevance to their weed biocontrol programme, and to Dr P. Harris (Agriculture Canada, Regina) for arranging this funding. These funds also supported two temporary technicians in Londo.n, A. D. Seccombe and R. D. Wild, who did many of the dissections. Mr Wild also carried out a project on the inter-population variability of Urophora quadrifasciata and he kindly allowed us to use the results of his unpublished work in this paper. 

Plant nomenclature follows Moore et al. (1976) for European taxa, Feinbrun-Dothan (1978) for Israel and Davis (1975) for Turkey. The large genus Centaurea is divided into the subgenera proposed by J. Dost61 (in Moore etaf., 1976) . Host records that have not been confirmed by the examination of reared material are followed by a note or reference in square brackets. Unconfirmed records of doubtful validity are not included. Single rearing records that do not fit the normal host range of a species are marked 'aberrant host?' to draw attention to the possibility that they may only be hosts in a single area or the result of a few individuals of a large population ovipositing on the 'wrong' plant. [Frauenfeld, 1857,18631 maura [Frauenfeld, 1857,18631 maura [Frauenfeld, 1857,18631 ? maura maura [not reared] maura [Richter, 19701 angustifuscia [Freidberg, 1974bj hermonis Y solstitialis [Zwolfer, 1965; aberrant host ?] ?solstitialis [Zwolfer, 1965; aberrant host ?] stylata [Neuenschwander et [Kugler & Freidberg, 19751