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Stuttgart 1. August 1967 Nr. 177

Revision of the Phytomyza syngenesiae group (Diptera, Agromyzidae), including species hithertö known as Hr “Phytomyza atricornis Meigen” By Graham C.D. Griffiths, London JUL 9 1969 With 23 Figures

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I. Introduction

The small group of species treated in this paper includes two species, Phytomyza sSyngenesiae (Hardy) and P. horticola Goureau, which are of substantial economic importance. Hitherto these species have been confused in the literature, since they are not separable on the basis of external morphology. On present knowledge a reli- able diagnosis is possible only by examination of the male aedeagus. Dissection of the abdomen is necessary for this examination, except in the case of specimens whose aedeagus is extruded as during copulation. Since 1920 the above two species have generally been called „Phytomyza atricornis Meigen“, a name whose use I propose to discontinue for reasons which are explained below in the section headed “Notes on types and nomenclature”. An immense literature has accumulated on these two species. | have devoted my attention in this study mainly to taxonomic and biological

ı Papers, and have made no attempt to consider papers concerned primarily with control measures. } The figures of male genitalia in this paper are based on specimens which have been cleared in potassium hydroxide or sodium hydroxide, as is the practice in most contemporary work on the Agromyzidae. The descriptions given in this paper refer solely to characters which seem to me useful for purposes of identification or relevent to the classification proposed. Addi- tional information on the external morphology of the adults can be obtained from the very detailed descriptions of “Phytomyza atricornis Meigen” given by Henpeı, 1934 (probably based both on P. syngenesiae [Hardy] and P. horticola Goureau) and Merıs (1935), whose paper clearly refers to P. horticola Goureau, as is shown by his figures of the male genitalia. Some details of the female genitalia of horticola are given by Meuıs (1935) and SasakawA (1961 a). Studies of the larval morphology and biology ' of horticola (as “atricornis”) have been published by Merıs (1935), Anmap and GupTA

(1941), Bunr (1953), Kuropa (1960) and Treman and SEHGAL (1963). However the | only biological accounts certainly referable to syngenesiae are those referring to America (of which the most detailed is that of SmuLyAn, 1914) and New Zealand (see } Wart, 1923 and Keısey, 1937). The results of the important British studies by CoHEn

N I

2 STUTTGARTER BEITRÄGE ZUR NATURKUNDE Nr. 177

(1936) and Hussey and Gurney (1962) require clarification, since the identity of the species concerned is not clear. At least the latter used material of two species (syn- genesiae and horticola) in their work, but the details of which species was used in each particular experiment cannot now be reconstructed. Fortunately Dr. Hussey is continuing his work on this subject, and it is to be hoped that he will be able to clarify his earlier results in due course. It is also not clear whether the larval descriptions

given by Arıen (1958) and oe MeıkrE (1926) relate to P. syngenesiae (Hardy) or

P. horticola Goureau.

Some of the technical terms of phylogenetic systematics used in this paper (such as “apomorph”’, “plesiomorph” etc.) may perhaps still be unfamiliar to some readers. My use of these terms is in accordance with the definitions proposed in the works of Hennıc. A convenient summary of his views has recently appeared in English (Hennig, 1965).

The following abbreviations are used to indicate the location of material recorded in this paper:

AM The Australian Museum, Sydney

BM British Museum (Natural History), London

CSIRO Commonwealth Scientific and Industrial Research Organization, Canberra

DEI Deutsches Entomologisches Institut, Eberswalde

GCDG The author’s personal collection

GCRI Glasshouse Crops Research Institute, Littlehampton, Sussex, England

HEL University Zoological Museum, Helsinki

ISZ Institut für Spezielle Zoologie und Zoologisches Museum der Humboldt-Universität, Berlin

KAS Mr. K. A. Spencer’s collection, London

KB Universitetets Zoologiske Museum, Copenhagen

LUND Zoological Institute, University of Lund

MNHN Museum National d’Histoire Naturelle, Paris

NMI National Museum of Ireland, Dublin

USNM U.S. National Museum, Washington

WIEN, Naturhistorisches Museum, Vienna

Other abbreviations used in recording material are:

ex. examples em. emerged leg. legit (i. e. collected)

I have stated the collector’s name, where known, for all material collected by workers other than myself.

II. The Phytomyza syngenesiae group

General Description

Adult

External Morphology: Head (fig. 1) with the orbits not or only narrowly project- ing above the eye in lateral view. Two upper orbital bristles (ors) of about equal length

present, but only a single strong lower orbital (ori) except sometimes in aragonensis. Third antennal segment short and rounded. Eye distinctly higher than long (contrast

Phytomyza plantaginis Robineau-Desvoidy), with only very sparse pubescence (contrast

P. nigra Meigen).

Acrostichal hairs sparse (at most six to seven present in farfarella and lindbergi) or completely absent (in aragonensis and most specimens of syngenesiae and horticola): intra-alar hairs very few, usually only one or two presuturals present (occasionally there is also a single postsutural, but usually this is absent).

Wing (fig. 2) with the second costal segment (between the apices of r, and 13,3) 11/2—2 times as long as the fourth segment (between r,,, and m;;,).

1967 GRIFFITHS, REVISION PHYTOMYZA SYNGENESIAE GROUP Nr. 177 /3

Colour: Frons entirely yellow or occasionally with the upper part of the orbits slightly darkened. Face yellow, with a variable degree of infuscation in the hollows below the antennae. Antennae largely dark, at most with the first and second segments yellow-brown. Thorax almost entirely matt grey, with only the wing base and dorsal

Fig. 1. Head in lateral view of Phytomyza syngenesiae (Hardy). (Scale 0.1 mm.) Fig. 2. Wing of Phytomyza syngenesiae (Hardy). (Scale 1 mm.)

margin of the mesopleuron yellow. Femora largely dark, with yellow apices. Basal cone of ovipositor (2?) entirely grey dusted (aragonensis), or grey dusted basally but

' with its apical half or third shining (farfarella, syngenesiae and horticola).

Size small (wing length’not exceeding 2.6 mm).

ö genitalia: Surstylus (fig. 20) with a small group of hairs at its tip, only parti- ally fused with the epandrium (9th tergite), delimited by a distinct suture on its outer side. Aedeagus (figs. 5, 7”—9, 12, 13, 16, 17 and 21) with the basiphallus consisting of two long sclerites which diverge apically; mesophallus forming a characteristic elongate tubule; distiphallus arising from the base of the mesophallus. Ejaculatory bulb (fig. 6, 11, 15 and 19) relatively small.

2 genitalia: Spermathecae minute and orbicular (see SasakAawA, 1961 a).

Larva

Mouthparts and cephalopharyngeal skeleton as in the majority of Phytomyza spp. Mandibles each with two alternating teeth, the right mandible being longer than the left. Spiracles (both front and hind) small, with 6—12 bulbs.

ZH)

3 Fig. 3. Puparium of Phytomyza syngenesiae (Hardy). (Scale 1 mm.)

Biology

The larvae form linear mines in the leaves of various herbaceous plants, pupating within the leaf in a “pupal blister”. The puparium (fig. 3) lies with its ventral surface "adjacent to the surface of the leaf, with its front spiracles projecting ventrally through the epidermis, £

|

4 STUTTGARTER BEITRÄGE ZUR NATURKUNDE Neizae

The Phytomyza syngenesiae group as here conceived is well characterised as a monophyletic group by the form of the aedeagus, particularly the characteristic elon- gate form of the mesophallus. This is an apomorph (derivative) character not shown by other species of Phytomyza. Other clearly apomorph features of the group are the reduced number of acrostichal and intra-alar hairs, and the low ratio of costal seg- ments 2 to 4. Each of these features however occurs in certain other species of Phyto- myza, and it is not yet possible to judge whether they characterise the syngenesiae group as such, or indicate its synapomorphy with other species. |

All adult males and some adult females of the species of the Phytomyza syn- genesiae group may be distinguished by the key given below. On present information | the females of syngenesiae and horticola cannot be reliably separated; and although many females of farfarella can be recognised by their relatively numerous acrostichal | hairs, their range of variation in this respect overlaps with that of syngenesiae. Con- siderable caution is thus needed in any attempt to identify isolated females, and it must be accepted that in many cases an identification to species will not be possible. Larvae and puparia were available to me for only three species, aragonensis, syn- genesiae and horticola; these do not seem to exhibit any characters suitable for indi- vidual diagnosis, though I have little doubt that future studies will reveal statistical |

differences.

Key to adults ofthe Phytomyza syngenesiae group

1 3rd antennal segment (fig. 4) with a fringe of conspicuous long white pubescence. Acrostichal hairs completely absent. Aedeagus (Ö) as fig.5 . P. aragonensis n.sp. |

— Pubescence of 3rd antennal segment short and inconspicuous (fig. 1) . ... 2

2 Trochanters and apex of the front coxae rather bright yellow. Six acrostichal hairs | (acr) present. Aedeagus (6) as figs. 7 and 8: sclerites of basiphallus becoming. broad and obscurely defined towards their apex; distiphallus weak; mesophallus slightly longer than in other species of this group. (Canary Isles)

P. lindbergi Spencer. — Trochanters and apex of the front coxae dark, or at most yellow-brown . . . 3

3 Aedeagus (Ö) as figs. 16—18 and 21: distiphallus strongly developed, divergent from its base; mesophallus bent upwards only at its apex. Acrostichal hairs usually completely absent . . .. .. P. horticola Goureau,

— Aedeagus (Ö) as figs. 9, 10 ai Ta ale more weakly developed, divergent only towards its apex; mesophallus with about its apical third bent' upwards Fi a ee i 4

4 Usually four to seven ee ee present, rarely no more en two. Aedeagus (5) as fig. 9, with the sclerites of the basiphallus becoming broad and obscurely defined towards their apex; the sac below the mesophallus is large and usually’ somewhat sclerotised along its hind margins . . . .. P. farfarella Hendel

— Acrostichals often completely absent, at most one ee en isolated hairs present. Aedeagus (6) as fig. 12, with the sclerites of the basiphallus more well defined towards their apex; the sac below the mesophallus is smaller and virtually mem- branous (easily distorted in preserved specimens) . . . P. syngenesiae (Hardy) |.

| | ! |

III. Treatment of species

Phytomyza aragonensis.n.sp. | Holotype & in the Institut für Spezielle Zoologie und Zoologisches Museum der Humboldt-Uni- versität, Berlin. | Adult Development of second lower orbital bristles (ori) variable (strongly developed, | about half the length of the first ori, in the holotype; vestigial in one of the females; \

1967 GRIFFITHS, REVISION PHYTOMYZA SYNGENESIAE GROUP Nr. 177 /5

strongly developed on one side only in the other female). 3rd antennal segment (fig. 4) with a fringe of conspicuous long white pubescence.

Acrostichal hairs (acr) completely absent.

Trochanters and apex of the front coxae yellow-brown or brown. Basal cone of ovipositor (2) entirely grey dusted.

Pr

Figs. 46. Phytomyza aragonensis n. sp. holotype &: 4, third antennal segment; 5, aedeagus in lateral view; 6, ejaculatory bulb. (Scale 0.1 mm.)

ö genitalia: Aedeagus (fig. 5) with the distiphallus only weakly divergent towards its apex; mesophallus with its apical third characteristically sinuate; the sac below the mesophallus is large (similar to that of farfarella), with its hind margins strongly sclerotised. Ejaculatory bulb (fig. 6) only weakly asymmetrical.

Material examined: Holotype 8; 2 22 paratypes, from larvae and puparia 2. V1. 33 in leaves of Lactuca tenerrima, Albarracin, Aragon, Spain, em. mid-V1. 33, leg. HErınG no. 4225 (ISZ).

The type series was recorded as “Phytomyza atricornis Meigen” by Herınc (1936). They were bred from narrow leaf-mines similar to those of syngenesiae and horticola. Pupation took place on the underside of the leaf.

It should be noted that the species with very long antennal pubescence occurring

‚on Lactuca in North America, Phytomyza lactuca Frost, is not referable to the syn- ‚genesiae group as defined in this paper; nor is the European P. ciliata Hendel (asso- eiated with Chrysanthemum leucanthemum). The resemblance of aragonensis to these species in respect of its long antennal pubescence is clearly to be interpreted as ‚convergence.

Phytomyza lindbergi Spencer Phytomyza lindbergi Spencer, 1957 (p. 1—2) and 1965 a. Holotype &, La Palma (Canary Isles), in the University Zoological Museum, Helsinki.

Adult

Only a single strong lower orbital bristle (ori) present (2nd ori vestigial). Antennal oubescence short.

Six acrostichal hairs (acr) present.

Trochanters and the apex of the front coxae rather bright yellow.

ö genitalia: Aedeagus (figs. 7 and 8) with the sclerites of the basiphallus becom- ‚ng broad and obscurely defined towards their apex; distiphallus weakly developed; ‚nesophallus slightly longer than in the other species of this group; the sac below the

I |

6 STUTTGARTER BEITRÄGE ZUR NATURKUNDE Nr. 177

8

Figs. 7—8. Phytomyza lindbergi Spencer holotype &: 7, aedeagus in lateral view; 8, aedeagus in ventral view. (Scale 0.1 mm.)

mesophallus is membranous and appears smaller than in the other species (the latter character however requires checking on fresh material). (Ejaculatory bulb lost.) (? unknown.)

Material examined: Holotype ö, El Paso (600 metres), La Palma, Canary Isles, 4.1V.50, leg. Linpeerc (HEL).

The holotype is still the only known specimen.

Spencer (1957) originally compared this species with Phytomyza asteris Hendel, but it is evident from the male genitalia that it should be included in the syngenesiae group. In asteris the aedeagus is of a somewhat different form, lacking the character- istic mesophallus of the syngenesiae group.

Phytomyza farfarella Hendel

Phytomyza farfarella Hendel, 1935 (p. 401). Holotype ®, Jugoslavia, in the Naturhistorisches

Museum, Vienna. Phytomyza atricornis Meigen sensu Griffiths, 1964 and earlier records referring to Iceland (nomen

dubium). Adult

Usually only a single lower orbital bristle (ori) present, but a vestigial 2nd (lower) ori is found in a few specimens (including the holotype). Antennal pubescence short.

Usually 4—7 acrostichal hairs (acr) present, but no more than two in some Icel- andic specimens.

Coxae and trochanters dark (contrast lindbergi). Basal cone of ovipositor (?) grey dusted basally, but with about its apical third shining.

ö genitalia: Aedeagus (figs. 9 and 10) with the sclerites of the basiphallus be | coming broad and obscurely defined towards their apex; distiphallus weakly developed, sometimes hardly visible; mesophallus as in syngenesiae, with about its apical third bent upwards; the sac below the mesophallus is larger than in syngenesiae and more | strongly sclerotised, especially along its hind margins. Ejaculatory bulb (fig. 11) usually | with a rather longer ° “stalk” than in the other species of this group.

Material examined:

Yugoslavia — 2 22 (one the holotype), Bleder See, V., leg. Henper (WIEN).

1967 GRIFFITHS, REVISION PHYTOMYZA SYNGENESIAE GROUP NETZ.

10

Figs. 9—11. Phytomyza farfarella Hendel &, Iceland: 9, aedeagus in lateral view; 10, distiphallus in dorsal view; 11, ejaculatory bulb. (Scale 0.1 mm.)

Germany — 1 Ö ex Taraxacum officinale agg., Berlin-Finkenkrug, em. 3. VI. 24, leg. Hering no. 2447 (ISZ). 2 86 1 2, same plant, Berlin-Frohnau, em. 24-27. V1. 25, leg. Herınc (ISZ). 25 ö, same plant and locality, em. 31. VII. 26, leg. HErına no. 2971 (ISZ). 1 Ö, same plant, Güntersberg an der Oder, em. 20. VI. 29, leg. Herınc no. 1066 (ISZ).

Denmark — 1 ? ex Leontodon autumnalis, Lemvig, em. 5. V1. 50, leg. Sonperur (ISZ).

Sweden — 1 ö ex Taraxacum sp., Hedlandet, Södermanland, em. 30. VI. 43, leg. Lunovist (LUND): 1 $, same host and locality, em 15. VII. 43, leg. Lunnovıst (LUND).

Finland — 1 Ö, Saltvik, Äland, VII. 42, leg. Frey (HEL).

Ireland — 1 ö, Lough Rask, near Ballyvaughan, Co. Clare, 3-8. VII. 60, leg. Cor (BM). 1 8 from the Harıpar collection (NM]).

Faroes — 4 86,2 2? from the islands of Streymoy, Nölsoy and Svinoy, as recorded in GRIFFITHsS, 1964 (KB).

Iceland — 44 ex. from localities in South and South-East Iceland, as recorded in GRIF- rırHs, 1964 (LUND and Göteborg Natural History Museum). 1 5, Skidastadalaug, Skagafjardarsysla (North Iceland), 27. VII. 33, leg. Tuxen (KB). 4 5 ö, Heimaey, Westmann Isles, 23. VII. 65 (2 ex.) and 13-14. VI. 66, leg. Anpersson (LUND).

Other Fennoscandian localities given in the literature are Paanajärvi, near Kuusamo in northern Finland (Frey, 1946) and Gotland (Fridhem), Sweden (Ryoen, 1952). This is the only species of the syngenesiae group occurring in Iceland and the Faroes. I failed to distinguish it from syngenesiae and horticola when writing my work , onthe Agromyzid fauna of these islands (Grırrırahs, 1964), and consequently some of the comments on “atricornis” in that paper require revision. Since there is no evidence that farfarella is a species likely to be spread by human agency, I think it must be re- garded as part of the indigenous fauna of these islands which is believed to have entered them during an interglacial period over a land-bridge from Europe. The spe- cies seems particularly common in the vicinity of hot springs in Iceland. My figure of the male genitalia of this species in that paper (Grırrıtus, 1964, fig. 4) was slightly misleading in one respect: the ejaculatory bulb figured is disproportionately large owing to a confusion in the scale of magnification. The ejaculatory bulb in this species is of similar relative size to that of other members of the syngenesiae group.

Phytomyza syngenesiae (Hardy) ‚ Chromatomyia Syngenesiae Hardy, 1849 (p. 391), in part. Neotype &, Ireland, by present desig- "nation in the British Museum (Natural History): type locality, Scotland.

8 STUTTGARTER BEITRÄGE ZUR NATURKUNDE Nr. 177

Phytomyza nigricornis M. Macquart sensu auctt, (e. g. Currıs, 1845) (nec Phytomyza nigricornis

M. Macquart, 1835). Phytomyza geniculata M. Macquart sensu auctt, (nomen dubium preoccupied by Phytomyza geni-

culata Brulle, 1832). Phytomyza lateralis Fallen sensu auctt. (e.g. GourEAu, 1851 and RosıneAu-Desvoipy, 1851) (nec

Phytomyza lateralis Fallen, 1823). Phytomyza albiceps Meigen sensu auctt. (e. g. KALtengAach, 1874 [in part] and Watt, 1923) (nec

Phytomyza albiceps Meigen, 1830). Phytomyza affinis Fallen sensu auctt, (e. g. French, 1900) (nec Phytomyza affinis Fallen, 1823). Phytomyza Chrysanthemi Kowarz in Lintner, 1891 (p, 242—243). Lectotpye &, U.S.A., by present designation in the Deutsches Entomologisches Institut, Eberswalde. Syn. nov. Phytomyza atricornis Meigen sensu Hendel, 1920 (in part) and widely in subsequent literature

(nomen dubium). Adult Only a single lower orbital bristle (ori) present. Antennal pubescence short. Acrostichal hairs (acr) often completely absent, at most one to four isolated hairs present. (Such specimens with a few acr present occur frequently in this species, but only rarely in horticola.) Basal cone of ovipositor (?) grey dusted basally, but with its apical half or third

shining.

12

Figs. 12—15. Phytomyza syngenesiae (Hardy) & ex Chrysanthemum vulgare, Ireland: 12, aedeagus in

lateral view; 13, aedeagus in ventral view; 14, distiphallus in dorsal view; 15, ejaculatory bulb. (Scale 0.1 mm.)

& genitalia: Aedeagus (figs. 12—14) with the sclerites of the basiphallus becom-

ing rather widely separated distally, more well defined towards their apex than in farfarella and lindbergi; distiphallus rather weakly developed, divergent only towards its apex; mesophallus with about its apical third bent upwards (as also in farfarella); the sac below the mesophallus is membranous (easily distorted in preserved specimens), rather smaller than in farfarella. Ejaculatory bulb (fig. 15) usually distinctly asym- metrical. Material identified by examination of the aedeagus (Öö only) Ireland — 1 Öö neotype (by present designation) ex Senecio jacobaea, Doolin, Clare, em. 30. VIII. 66, leg. Grirrıtus (deposited in the British Museum [Natural Hi- story]). 1 6 ex Senecio sp., Dalkey, em. VI. 04 (NMI]). 1 ö ex Sonchus arvensis, Poulsallagh, Clare, em. 20. V1.65 (GCDG). 1 ö ex Chrysanthemum leucanthemum,

Caherconnell, Clare, em. 26. VI. 65 (GCDG). 2 868 ex Chrysanthemum vulgare,

Burren (near Ballyvaughan), Clare, em. 16 and 22. IX. 66 (GCDG).

1967 GRIFFITHS, REVISION PHYTOMYZA SYNGENESIAE GROUP Nr. 177 /9

Great Britain

Scotland — 1 ö ex Chrysanthemum argyrophyllum, Edinburgh, em. 15. VI. 53, leg. SPENCER (KAS).

Wales — 1 ö ex Inula crithmoides, Burry Holms, Gower, em. 7. X. 62 (GCDG).

England — 1 Ö ex Senecio cruentus, Bristol, em. IV. 20, leg. A. L. Sumu (BM).1 & ex Chrysanthemum sp., Putney, London, em. VI. 18, leg. Cant (BM). 7 68 & Chrysanthemum sp. (cultivated), Littlehampton, Sussex, leg. Hussey and GurnEY (GCRI). 1 ö ex Chrysanthemum sp. (cultivated), Kensington, London, em. 1. V1l. 15, leg. Rosıson (BM). 1 Ö ex Serratula tinctoria, Woodside Park, London, em. 10. VIII. 53 (GCDG). 1 ö ex Sonchus asper, Whitby, Yorks., em. 15. VII. 37, leg. Brırrrn (BM). 1 ö ex Arnoseris minima (labelled “Lapsana pusilla”), Middlesex, em. 3. VII. 23, leg. Hraru (BM).

France — 1 ö ex Anthemis sp., Asnieres, em. 7. 1V. 1889, leg. Lesne (MNHN). 1 6 ex Chrysanthemum leucanthemum, 1899, leg. BLancnarp (MNHN). 1 ö ex Chry- santhemum sp., Paris, em. 14.V1I.46 (MNHN). 1 & ex Inula crithmoides, La Baule, Loire inf., em. 15. IV. 43, leg. Bunr no. 4965 (152).

Denmark — 1 ö ex Daucus carota, Copenhagen, em. 18. IX. 29, leg. Gupman (ISZ). 1 © ex Senecio squalidus, Copenhagen Botanical Gardens, em. 15. XII. 65, leg. SorENSsEN (GCDG).

Sweden — 1 Ö ex Carduus crispus, Hälsingborg, Skäne, em. 3. IX. 23, leg. Ryotn (LUND). 1 5 ex Taraxacum sp., Hälsingborg, Skäne, em. 18. V11.23, leg. Ryoen (LUND).

‘ Finland — 1 ö ex Chrysanthemum sp., Hoplax, near Helsinki, 1936 (HEL).

Germany — 1 Ö ex Othonna integrifolia, Berlin Botanical Gardens, em. 5. III. 34, leg. Herınc (ISZ). 1 ö ex Xeranthemum annuum, Berlin, em. 28. VII. 24, leg. Herınc no. 2584 (ISZ).

Yugoslavia — 1 ö ex Sonchus oleraceus, Lesina, Hvar Island, em. IV. 29, leg. Bunr (ISZ).

Spain — 1 ö ex Sonchus oleraceus, Irun, em. 10. IV. 33, leg. Herınc no. 4101 (ISZ).

Canary Isles — 1 Ö ex unidentified plant, Los Arucas, Gran Canaria, em. 21. II. 63, leg. Spencer (KAS). 1 6 (caught), Maspalomas, Gran Canaria, 9-10. III. 50, leg. Linpserc (HEL). 1 ö (caught), Pico de Teyde, Tenerife, 21-22. VII. 31, leg. Frey (HEL). 1 ö (caught), La Llanos, La Palma, 8. VIII. 31, leg. StorA (HEL).

"Canada — 1 ö ex Crepis sp., Edmonton, Alberta, em. 4. VI. 66, leg. Senucar (in Mr. SeHcaAr’s collection). 1 Ö ex Senecio sp., Frank, South Alberta (4,000 ft.), em. 1. VII. 66, leg. Spencer (KAS).

U.S.A. — 2 ö6 syntypes of Phytomyza chrysanthemi Kowarz (one of which is hereby designated lectotype), presumably Boston, Massachusetts (DEI): host - plant not stated, but presumably a cultivated Chrysanthemum species. 1 ö ex “daisy”, Glen- dale, Rhode Island, em. 16. V. 10 (USNM). 1 ö ex “marguerite”, Kingston, Rhode Island, em. 6. I. 16, leg. Stone (USNM). 1 ö ex lettuce (Lactuca sp.) in greenhouses, Puyallup, Pierce Co., Washington, em. 1. 1. 53, leg. Frıck (USNM). 1 ö ex Sonchus sp., Seattle, Washington, em. IV. 41 (USNM). 1 6 ex Senecio eruentus, San Francisco, California, em. 1. VI. 65, leg. Toscnı & TAuser (BM). 1 5 ex Dahlia sp., San Francisco, California, leg. Wuıtney (USNM). 1 & ex Chry- santhemum sp., Alameda Co., California, em. 21. IV. 11, leg. Arprıch (USNM). 1 8 ex Zinnia elegans, Albany, Alameda Co., em. 27. IX. 48, leg. Frick (USNM). 1 ö ex Picris echioides, Albany, Alameda Co., California, em. 3. VIII. 48, leg. Frick (USNM). 1 & ex Lactuca sp., Albany, Alameda Co., California, em. 21. XI. 31, leg. Frick (USNM). 1 & ex Matricaria sp., Berkeley, Alameda Co., California,

‚_ em. 18. VI. 48, leg. Frıck (USNM). 1 ö ex Sonchus asper, Berkeley, Alameda Co.,

10 STUTTGARTER BEITRÄGE ZUR NATURKUNDE Ne. 1772

California, em. 21. VII. 48, leg. Frıck (USNM). 1 ö ex Sonchus oleraceus, Ber- | keley, Alameda Co., California, em. 1. VIII. 65, leg. ToscHı and TAuser (KAS). 1 ö ex Silybum marianum, Berkeley, Alameda Co., California, em. 23. XII. 48, | leg. Frıck (USNM). 1 ö ex Pisum sativum, Berkeley, Alameda Co., California, em. 14. VII. 17, leg. Arorıch (USNM). 1 ö ex C'ynara scolymus, Half Moon Bay, California, em. 11. V. 37, leg. Lance (USNM), 1 ö ex Helianthus californicus, ' ? California, leg. Wuırney (USNM). Australia — 1 Ö ex Senecio jacobaea, Leongatha, Victoria, em. 16. VII. 27, leg. Huı (CSIRO). 2 8& (caught), same locality, undated and 31. VIII. 27, leg. Hu (CSIRO). 1 ö (caught), Bronte, New South Wales, 8. IX. 54, leg. McAıpıne (AM). 1 ö (caught), Belltrees, near Scone, New South Wales, 19. X. 56, leg. McAıpine (AM). 1 ö (caught), North Beach, Bellinger River, New South Wales, 16. XI. 64, leg. McAırpıne (AM).. 1 ö (caught), Sydney University, New South Wales, 23. IX. 54, leg. McAıpine (AM). New Zealand — 2 && ex Sonchus sp., Canterbury, em. 11. VII. 63, leg. Yoons (GCDG).

The absence of records of this species from the Eastern part of the Palaearctic region hardly seems significant in view of the limited material of the syngenesiae group available to me from that area. I think it probable that syngenesiae is a holarctic spe- cies which has been able to spread across the former land connection between Siberia and Alaska. Its extensive distribution in North America suggests that it is not a recent . introduction there.

I have no doubt that the occurrence of this species in Australia and New Zealand | is the result of recent introduction. There seems no significant morphological differen- tiation in the material I have seen from these countries, and the recorded host range there includes many introduced plants. |

It is evident from the records given above that this species usually develops in Compositae hosts. The range of Compositae attacked is clearly very wide, although there are no records for a few genera (e.g. Aster and Solidago). Whether these apparent gaps are fortuitous or represent real discontinuities in the host range is not yet clear.

The occurrence of this species on hosts other than Compositae appears to be rare, and may be regarded as xenophagy. Details of one North American specimen labelled as bred from Pisum (Leguminosae) have been given above, and Frıck (1959) also records Mentha sp. and Stachys bullata (Labiatae), Malva borealis (Malvaceae) and E | Melılotus indica (Leguminosae) as hosts of “Phytomyza atricornis Meigen” in North . America. It seems probable that some or all of these records refer to syngenesiae (since horticola is not known in North America), but confirmation from the dissection of males bred from these hosts seems desirable before they can be finally accepted. From I Australia I have received one female presumably referable to this species which was | bred from Urtica incisa (Urticaceae) (Jenolan Caves, 2,000 ft., New South Wales, . leg. Wıeuro [AM]), and Henper (1934) also recorded a specimen of “atricornis” bred from Urtica ferox in New Zealand. Warr’s (1923) and Harrıson’s (1959) lists of hosts of “atricornis’ in New Zealand also include Plantago spp. (Plantaginaceae) and Melilotus sp. (Leguminosae). However the former record requires careful checking in view of the possibility of confusion with Phytomyza plantaginis Robineau-Desvoidy; and the latter record was clearly doubtful, as Watt prefixed it with the word “possibly”. From Europe I have discovered only one case of syngenesiae having been bred from a non-Compositae host, the record for Daucus (Umbelliferae) given above. But it would be unwise to assume from this evidence that the host tolerance of European strains of syngenesiae is less than that shown in other parts of the species’ range. The abundance | of horticola on non-Compositae hosts must inevitably tend to mask the rarer occurrence N of syngenesiae on such hosts in areas where both species occur. Any apparent diffe-

1967 GRIFFITHS, REVISION PHYTOMYZA SYNGENESIAE GROUP Nest

rence in host tolerance in the Palaearctic region should therefore be regarded with caution unless confirmed by laboratory experiments.

Complete lists of the host records which seem attributable to this species (subject to the remarks above) in areas where no other species of the syngenesiae group occurs have been given by Spencer (1963) for Australia, HAarrıson (1959) for New Zealand and Frıck (1959) for North America. One correction of Frıck’s list should however be noted. The series which he bred from Gnaphalium leucocephalum in California does not represent a species. of the syngenesiae group, but a species related to the European Phytomyza ciliata Hendel and P. farfarae Hendel.

Phytomyza horticola Goureau

Phytomyza horticola Goureau, 1851 (p. 148--149). Neotype &, Germany, by present designation in the Institut für Spezielle Zoologie und Zoologisches Museum der Humboldt-Universität, Berlin: type locality, France.

Phytomyza geniculata M. Macquart sensu auctt, (e. g. RosıneAu-Desvoıpy, 1851 and GouRrEAU, 1861 and 1869) (nomen dubium preoccupied by Phytomyza geniculata Brulle, 1832).

Phytomyza cucumidis J. Macquart, 1854 (p. 235) (nomen nudum). Syn. nov.

Phytomyza Tropaeoli Dufour, 1857 (p. 45). Types lost: type locality, France. Syn. nov.

Phytomyza Fediae Kaltenbach, 1860 (p. 250). Types lost: type locality, Germany. Syn. nov.

Phytomyza Linariae Kaltenbach, 1862 (p. 83) and 1874. Types lost: type locality, Germany. Syn. nov.

Phytomyza Pisi Kaltenbach, 1864 (p. 305) and 1874. Types lost: type locality, Germany. Syn. nov.

Phytomyza albiceps Meigen sensu auctt. (e. g. KALTEnBAcH, 1874, in part) (nec Phytomyza albiceps Meigen, 1830). ;

Phytomyza subaffinis Malloch, 1914 (p. 335). Holotype ©, Formosa, in the Hungarian Natural History Museum, Budapest. Syn. nov.

Phytomyza atricornis Meigen sensu Hendel, 1920 (in part) and widely in subsequent literature (nomen dubium).

16

17

IF igs. 16—20. Phytomyza horticola Goureau & ex Linaria, Italy: 16, aedeagus in lateral view; ‚17, aedeagus in ventral view; 18, distiphallus in dorsal view; 19, ejaculatory bulb; 20, surstylus and

part of 9th tergite (epandrium) in posteroventral view. (Scale 0.1 mm.)

12 STUTTGARTER BEITRÄGE ZUR NATURKUNDE Nr. 177

Napomyza lactucae Vimmer, 1926 (p. 117). Syntype 9, Czechoslovakia, in Mr. K. A. SPpEncER’s

collection, London. Syn. nov. Phytomyza bidensivora Seguy, 1951 (p. 313—314), Pauusan, 1953 and Spencer, 1959. Lectotype &, Madagascar, by present designation in the Museum National d’Histoire Naturelle, Paris. Syn. nov.

Adult

Only a single lower orbital bristle (ori) present. Antennal pubescence short.

Acrostichal hairs (acr) usually completely absent, rarely one or two isolated hairs present.

Basal cone of ovipositor (?)grey dusted basally, but with its apical half or third shining.

ö genitalia!: Aedeagus (figs. 16—18) with the distiphallus strongly developed, divergent from its base; mesophallus more evenly curved than in syngenesiae and farfarella, bent upwards only at its apex; the sides of the sac below the mesophallus form a pair of sclerotised plates which unite apically (fig. 17); these are largely trans- parent but with a variable degree of pigmentation along their margins. Ejaculatory

bulb (fig. 19) usually almost symmetrical, rarely as strongly asymmetrical as that figured _

for syngenesiae (compare fig. 15).

Materialidentified by examination of the aedeagus (d& only)

Ireland — 1 ö ex Pisum sativum, from the Hauıpar collection (NM]).

England — 1 ö ex Vicia sepium, Faversham, Kent, em. 7. V1l. 54 (GCDG). 2 66 ex Dahlia sp., Littlehampton, Sussex, leg. Hussey & Gurney (GCRI). 1 d ex Allium sativum in my garden at Barnet, London, em. 24. VI. 66 (GCDG).

France — 1 ö ex Lepidium draba, Hyeres, em. 21. V. 35, leg. Wırson (BM). 1 ö ex

Lycium sp., em. VII. 26, leg. Lesne (MNHN). 1 ö ex Senecio vulgaris, Tence (H.L.), em. 8. VII. 27, leg. MenevaL (MNHN). 1 ö ex Arctium lappa, Paris, em.

28. V1. 53, leg. Spencer (KAS). 1 Ö ex Cirsium vulgare, La Baule, Loire inf., em. 5. VI. 43, leg. Bunr no. 386 (ISZ). 1 ö ex Allium cepa, Cherbourg, em. 10. VI. 28, leg. Herınc (ISZ).

Switzerland — 1 Ö ex Inula hirta, Lenzerheide, em. 16. VII. 25, leg. Hopp (ISZ).

Denmark — 1 Ö ex Taraxacum officinale agg., Lemvig, Denmark, em. 10. X. 53, leg. SONDERUP (KAS).

Sweden — 1 ö ex Brassica campestris var. capitata, Hälsingborg, Skäne, em. 19. VIII. 23, leg. Ryoen (LUND). 1 ö ex Linum usitatissimum, Hälsingborg, Skäne, em.

2. IX. 24, leg. Rypen (ISZ). 1 6 ex Pisum sativum, Hedlandet, Södermanland,

em. 23. VII. 43, leg. Lunnpovist (LUND). 1 ö ex Melilotus albus, Hälsingborg, |

Skäne, em. 2. VIII. 27, leg. Ryotn (LUND). 1 ö ex Matricaria sp., SONDERUP, em.

17. VII. 52, leg. Rroen (LUND). 1 ö ex Senecio vulgaris, Hälsingborg, Skäne,

em. 30. VI. 23, leg. Ryotn (LUND). 1 ö ex Helichrysum sp., Hedlandet, Söder- |

manland, em. 22. VIII. 43, leg. Lunpoviıst (LUND). Finland — 1 ö (caught), Forvas, leg. Frey (HEL).

Germany — 1 Ö ex Adonis aestivalis, Erfurt, Schwellenburg, Thuringia, em. 12. VI. | 58, leg. Bunr no. 1352 (KAS). 1 6 ex Brassica napus, Kiel, em. IX. 28, leg.

Bıunck (ISZ). 1 ö ex Conringia orientalis, Rostock Botanical Gardens, Mecklen- burg, em. 5. VII. 49, leg. Bunr no. 182 (ISZ). 1 ö ex Thlaspi arvense, Masser- berg, Thuringia, em. 11. X. 58, leg. Bunr no. 1449 (ISZ). 1 ö ex Papaver sp., Jerichow (Elbe), em. 8. VII. 23, leg. Herınc no. 2305 (ISZ). 1 ö ex Papaver sp., Aschersleben, leg. Norte, V1.50 (DEI). 1 ö ex Cleome dodecandra, Rostock, Mecklenburg, em. 27. VII. 32, leg. Bunr (ISZ). 1 ö neotype (by present designa- tion) ex Tropaeolum majus, Jerichow (Elbe), em. 23. VII. 22, leg. Hering no. 2085 (ISZ). 1 & ex Cerastium semidecandrum, Berlin-Frohnau, em. 13. VI. 26, leg.

! The description relates to typical material from the Palaearctic and Oriental Regions. Geographi- cal variation in the Ethiopian Region is discussed below,

1967 GRIFFITHS, REVISION PHYTOMYZA SYNGENESIAE GROUP Nr. 177/13

Herne (ISZ). 1 ö ex Salicornia herbacea, Rostock Botanical Gardens, Mecklen- burg, em. 12. VIII. 37, leg. Bunr (ISZ). 1 ö ex Ononis hircina, Rostock, Mecklen- burg, em. 26. VI. 35, leg. Bunr (ISZ). 1 Ö ex Pisum sativum, Aschersleben, leg. Norte, V1. 50 (DEI). 1 ö ex Pimpinella anisum, Rostock Botanical Gardens, Meck- lenburg, em. 12. VII. 36, leg. Bunr (ISZ). 1 ö ex Tordylium persicum, Mecklen- burg, em. 21. VI. 37, leg. Bunr (ISZ). 1 ö ex Asperugo procumbens, Schwäbischer Jura, em. 12. VIII. 23, leg. Hrrınc no. 2369 (ISZ). 1 ö ex Lithospermum offiei- nale, Bellinchen Mark, em. 4. VI. 26, leg. Herınc no. 2887 (ISZ). 1 6 ex Sym- phytum officinale, Kirschberge, Mühlhausen, Thuringia, em. 7. VI. 66, leg. Bunr no. 2800 (GCDG). 1 ö ex Echium vulgare, Bredow bei Nauen, em. 29. VII. 23, leg. Herınc no. 2337 (ISZ). 1 ö ex Satureia hortensis, Berlin-Frohnau, em. 3. IX. 26, leg. Herınc no. 2952 (ISZ). 1 6 ex Phlox drummondi, Rostock, Mecklenburg, em. 3. VIII. 35, leg. Bunr no. 77/7 (ISZ). 1 & ex Linaria vulgaris, Jerichow (Elbe), em. 9. VII. 23, leg. Herınc no. 2303 (ISZ). 1 d ex Valeriana sambucifolia, Rostock, Mecklenburg, em. 6. VI. 35, leg. Butir no. 682 a (ISZ). 1 ö ex Valeriana officinrlis, Jerichow (Elbe), em. 5. VII. 23, leg. Herınc no. 2302 (ISZ). 16 ex Cen- taurea jacea, Berlin-Dahlem, em. 27. VI. 42, leg. Herınc no. 4838 (ISZ). 1 ö ex Doronicum austriacum, Berlin Botanical Gardens, em. 24. VI. 23, leg. Herınd no. 2235 (ISZ). 1 ö ex Chrysanthemum corymbosum, Berlin Botanical Gardens, em. 28. VI. 23, leg. Herınc no. 2241 (ISZ). 1 ö ex Chrysanthemum vulgare, Ber- lin-Frohnau, em. 6. VII. 26, leg. Herınc no. 2946 (ISZ). 1 ö ex Ärtemisia dra- cunculus, Berlin-Frohnau, em. 13. VII. 26, leg. Herınc no. 2959 (ISZ). 1 d ex Senecio vernalis, Berlin, em. VI. 26, leg. Hope (ISZ). 1 6 ex Senecio vulgaris, Berlin-Dahlem, em. 12. VI. 48, leg. Herıns no. 5420a (ISZ). 1 ö ex Erigeron canadense, Berlin-Frohnau, em. 8. VI. 26, leg. Herına no. 2921 (ISZ). 1 d «x Anthemis tinctoria, Stempeda, Harz, em. 25. VI. 27, leg. Herınc no. 3158 (ISZ). 1 ö ex Taraxacum officinale agg., Berlin-Frohnau, em. 1. VIII. 26, leg. Herınc Bor 2971. USZ).

Austria — 1 Ö ex Centaurea jacea, Linz, Upper Austria, em. 1. VII. 62, leg. Herınd no. 6792 (KAS).

Poland — 1 ö ex Hesperis matronalis, Rabendorf, Silesia, leg. Teıcaer (ISZ).

Russia — 1 ö ex Brassica juncea, Rostov-on-Don, em. 17. VI. 27, leg. Mamonow (ISZ). 1 8 ex Ononis hircina, Rostov-on-Don, em. 12. VII. 27, leg. Mamonow (ISZ). 1 S ex Cicer arietinum, Rostov-on-Don, em. 7. VII. 27, leg. MAmonow ne 1 8 ex Vigna sinensis, Rostov-on-Don, em. 28. VI. 27, leg. Mamonow ISZ).

Yugoslavia — 1 Ö ex Chrysanthemum coronarium, Jelsa, Hvar Island, em. 12. V. 65,

leg. Herınc no. 7326 (ISZ). 1 6 ex Cirsium creticum, Jelsa, Hvar Island, em. 27. V.65, leg. Herınc no. 7399 (ISZ).

Italy — 3 86 ex Linaria sp., Ostia, near Rome, em. 18. IV. 54 (GCDG). 1 d

Senecio doria, Rome, em. 7. 1. 65, leg. Frıck (KAS).

Corsica — 1 Ö ex Isatis sp., Corte, em. IX. 33, leg. Bunr (ISZ).

Sicily — 1 Ö (caught), Sinofsi, Mount Etna, 8. IV. 64, leg. Spencer (KAS).

Spain — 2 Öd ex Sisymbrium orientale, Albarracin, Aragon, em. VI. 33, leg. Herına no. 4222 (KAS and ISZ). 3 &Ö (caught), Mongat, near Barcelona, 21. II. 10, coll. Lichtwarpr (DEI).

Portugal — 1 d ex Gazania sp., Lisbon, em. 4. IV. 53, leg. Spencer (KAS).

‚ Azores — S. Miguel. 1 ö, Ponta Delgada, 12. V. 38, leg. StorA (HEL). 1 Ö, Furnas,

19-21. V. 38, leg. Stori (HEL). Terceira. 1 8, Santa Barbara, 7. VI. 38, leg. Frey (HEL). 1 6, Achada, 3-4. VI.

‚38, leg. Frey (HEIL).

| Pico. 1 ö, Madalena, 6-9. VII. 38, leg. Stork (HEL).

14 STUTTGARTER BEITRÄGE ZUR NATURKUNDE Nr. 177

Fayal. 3 öÖ, Caldeira, 4. VII. 38, leg. Frey (HEL).

Flores. 1 ö, Santa Cruz, VI. 38, leg. Storä (HEL). 1 ö, Ribeira Fazenda, VI. 38, leg. StorA (HEL).

Corvo. 1 ö, Caldeiras, 24. VI. 38, leg. Storä (HEL).

Madeira — 1 Ö, Funchal, 1-8. V. 38, leg. Frey (HEL). 1 Ö, Porto Novo, 5. V. 38, leg. Frey (HEL).

Canary Isles — 2 ÖÖ ex Malva neglecta, Santa Cruz, Tenerife, em. 16-17. Ill. 26, leg. Herınc no. 2765 (ISZ). 2 66 ex Lathyrus tuberosus, Laguna, Tenerife, em. 14-15. III. 26, leg. Herınc no. 2778 (KAS and ISZ). 1 ö ex Plantago lanceolata, Puerto de Cabras, Fuerteventura, em. 21. III. 26, leg. Herınc no. 2784 (ISZ). 1 6 ex Bystropogon plumosum, El Paso, Palma, em. 11. IV. 26, leg. Herınc no. 2830 (ISZ). 1 8 ex Lithospermum arvense, Puerto de Cabras, Fuerteventura, em. 27. III. 26, leg. Herınc no. 2788 (ISZ). 1 ö ex Bidens pilosus, Santa Cruz, La Palma, em. 16. IV. 26, leg Herınc no. 2808 (ISZ). 1 ö ex Carduus baeocepha- lus, Laguna, Tenerife, em. 28. IV. 26, leg. Hrrınc no. 2875 (ISZ). 2 56 (caught), Las Palmas, Gran Canaria, 28-30. V1. 31, leg. Frey and Stori (HEL). 1 ö (caught), Chilegua, Fuerteventura, 4-14. Ill. 49, leg. Linpsere (HEL). 2 6& (caught), Haria, Lanzarote, 19. III. 49, leg. Liınpeers (HEL).

Morocco — 1 ö ex Compositae sp., Casablanca, em. 28. I. 66, leg. Spencer (KAS). Egypt — 1 Ö ex Pisum sativum, Heliopolis, em. 20. XII. 21, leg. Harcreaves (BM). 1 ö ex Solanum melongena, Alexandria, em. 29. Ill. 56, leg. Hammao (1SZ). Palestine — 1 ö ex Eruca sativa, Tel Aviv, leg. BoDEnHEIMER (ISZ). 1 Ö& ex Papaver somniferum, Tel Aviv, leg. Bopenneimer (ISZ). 1 Ö ex Pisum sativum, Tel Aviv, leg. BopenhEImer (ISZ). 1 Ö ex Linaria vulgaris, Tel Aviv, em. 21-30. IV. 26, leg. BopenHEmMER (ISZ). 1 6 ex Chrysanthemum sp., Tel Aviv, leg. BoDENHEIMER

(ISZ). 1 ö ex Sonchus oleraceus, Tel Aviv, em. II. 26, leg. BopEnHEImEr (ISZ).

India — 1 ö ex Melilotus parviflora, Lyallpur, Punjab, em. 19. II. 23, leg. BAHADuRr (BM). 1 6 ex Pisum sativum, Lyallpur, Punjab, em. 19. III. 20, leg. BAHADUR (BM). 1 ö ex Chrysanthemum sp., Lyallpur, Punjab, em. 16. III. 23, leg. Bana- pur (BM). 1 ö ex Sonchus asper, Lyallpur, Punjab, em. 23. III. 20, leg. BanApur (BM).

China — 1 ö ex Brassica f. oleifera, Shanghai, em. V. 33, leg. Horne (ISZ). 1 ö ex Papaver somniferum, Shanghai, em. V. 33, leg. Horne (ISZ). 1 6 ex Pisum sati- vum, Canton, leg. Horrman (BM). 1 8 ex Linum usitatissimum, Canton, leg. Horr- MAN (BM). 2 56 ex Carduus sp., Charbin, Manchuria, em. 5-10. VI. 51, leg. Arın (ISZ). 1 ö ex Helianthus annuus, Charbin, Manchuria, em. 8. V. 51, leg. Arın (ISZ).

Formosa — 1 Ö (paratype of Phytomyza subaffinis Malloch), Tainan, II. 1909, leg. SAuTER (Hungarian Natural History Museum). 1 ö ex Solanum indicum, Taipei, em. 7. IV. 65, leg. SasakawA (GCDG).

Japan — 1 ö ex Artemisia vulgaris, Ashoro, Hokkaido, em. 16. IV. 54, leg. SasA- KAwA (GCDG). 1 6 ex Nasturtium indicum, Kibune (Yamashiro), Honshü, em. 19. V. 53, leg. Sasakawa (GCDG). 1 & ex Chrysanthemum sp., Kibune (Yama- shiro), Honshü, em. 7.V.53, leg. SasakawAa (GCDG). 1 6