GALL MIDGES ECONOMIC IMPORTANCE - KrishiKosh

GALL MIDGESOFECONOMICIMPORTANCEH. F. BARNESM.A. (Oxford), Ph.D. (London)Rothamsted Experimental Station, HarpendenVOL. I:GALL MIDGES OF ROOT AND VEGETABLE CROPSWith a Foreword byC. T. GIMINGHAM. O.B.E.. B.Sc, F.R.I.C.Director, Plant Pathology Laboratory,Ministry of Agriculture and Fisheries.r^^6^^LONDONCROSBY LOCKWOOD & SON LTD20, TUDOR STREET. LONDON, E.C.41946

IN GRATEFUL MEMORY OFF. V. THEOBALDF. W. EDWARDSE. P. FELT

ForewordT, he small flies known as Gall Midges are ofmuch interest and importance. They include groups of speciesshowing the greatest diversity of habit, but the majority feedon plants in the larval stage and among these there are verymany that attack cultivated crops of many kinds, often causinginjury and serious losses. The Hessian Fly, sometimes responsiblefor much damage to wheat crops in America and elsewhere,is perhaps the best known, but there are many others—e.g. theSwede Midge, the Pear Midge, the Clover Seed Midge, theChrysanthemum Midge—that are exceedingly harmful insects.In spite of their biological interest and economic importance,the Gall Midges as a group have scarcely received from entomologiststhe attention that they deserve, although a few of theinjurious species have been intensively studied. Much of theliterature on the subject is difficult of access, and though thereare monographs describing and classifying the species of GallMidges, a reference book containing the biological andeconomic information available about these pests has beenlacking.Dr. Barnes has set himself the task of writing a comprehensiveaccount of all those species of Gall "Midges, throughout theworld, that are of economic interest either as pests of crops oras beneficial insects. He is well qualified to do so. Trained inapplied entomology under Professor F. V. Theobald, and intaxonomic work under Dr. F. W. Edwards, the author madea special study of the Gall Midges with Dr. E. P. Felt, thedistinguished American authority on this group of insects, andby his careful investigations during the past twenty years hashimself added much to our knowledge of them. While the5

6 Gall Midges of Economic Importanceemphasis in his work is on the bionomics of the insects anddoes not aim primarily at providing detailed descriptions, theinformation on the biology and habits of the species of economicimportance should enable entomologists to identify them.• The injurious species of Gall Midges present specially difficyltproblems for the economic entomologist because direct methodsof control are seldom, practicable, and suitable preventivecultural methods have to be sought. While it is true thatdetailed knowledge of the biology and life history of all insectpests is a prime requisite for the discovery of satisfactory meansof control, this applies with special force to pests that have tobe dealt with by cultural methods. To devise modifications infarming or gardening practice that will successfully enable thecrop to escape or withstand attack demands as full an understandingas possible of the relationship between the insectsconcerned and their plant hosts.Ecological studies on these lines have been carried out withseveral species of Gall Midges, work to which Dr. Barnes hasmade valuable contributions, but, as he points out, much stillremains to be done, and in collecting the available facts the igaps in our knowledge become apparent. The work, of whichthis^volume is the first part, is welcome both because it providesentomologists with a ready means of reference to the informationalready acquired, and because it will surely stimulateinterest in and attract workers to the further study of animportant but rather neglected group of insects.HARPENDEN.January, 1945.C. T. GIMINGHAM.

PrefaceT he gall midges ^ ^ _large family of small flies occurring throughout the world.The larvae of a great number are responsible for galls andother malformations on a large range of plants, while othersare zoophagous, feeding on aphides, coccids or scale-insects,white-flies and mites. Still others feed on rusts, mildews, etc.,and others are saprophytic (9).*Their galls have been described and recorded in manyscattered papers, and also in such very useful compilations asthose of Houard (85, 87, 88), Ross & Hedicke (158) andFelt (65, 70).Descriptions and the classification of the gall midges themselveshave similarly formed the subject of numerous papers,both monographic and otherwise, in many scientific journalsby such world-famous specialists as Kieffer, Riibsaamen andTavares in Europe, Felt in America and Mani in India.Comprehensive monographs also have been written byKieffer (103) and Rtibsaamen & Hedicke (178).In spite of the fact that several gall midges have gainedworld-wide notoriety as pests, and although there is muchscattered information in the literature concerning them as pests,there is nevertheless no comprehensive work on the gall midgesthat are of economic importance.The need for such a work of reference has long been felt, andthe reception accorded to my Material for a Monograph of theBritish Cecidomyidae or Gall Midges—British Gall Midges ofEconomic Importance I-V (14), besides making this very clear,gave me the incentive and encouragement to undertake the* Figures in brackets refer to the list of References on pages 83-93.7

8 Gall Midges of Economic Importanceambitious task of trying to fill this gap. It is believed that thepresent work will serve its primary purpose of making readilyavailable what is known concerning the gall midges of economicimportance throughout the world. In addition, it containsmuch thst should be of interest to the general biologist.It is based on a study of the literature, and on personalinvestigations on numerous species, carried out during the pasttwenty years. These investigations have involved more than150 species and necessitated breeding many thousands ofspecimens as well as the formation of an extensive collectionwhich now contains examples of most of the species of economicimportance.It is proposed to publish the work in separate volumes,each complete in itself. Vol. I will be Gall Midges of Root andVegetable Crops; Vol. II, Gall Midges of Fodder Cro^s; Vol. Ill,Gall Midges of Fruit; Vol. IV, Gall Midges of Ornamental Plantsand Shrubs. Subsequent volumes will deal with the gall midgesof trees, cereals, miscellaneous crops such as beverage plantsand herbs (culinary, aromatic and medicinal), and gall midgesthat are zoophagous, fungivorous and that attack weeds.Each volume will start, after a brief introduction, with a listof the crops (or prey in the case of the zoophagous midges) tobe dealt with, indicating the midges that attack them, the partof the plant attacked and the pages on which the informationconcerning the species can be found. These lists should facilitatethe recognition of the pests.The midges will usually be dealt with under the plants oranimals they attack, arranged alphabetically. The informationconcerning each will be presented in much the same order.Wherever possible this information is subdivided under sideheadings,but throughout the work stress is laid on the biologyand control. Thus, with comparatively well-known pests, immediatelyafter the name of the species there is a brief diagnosticcharacter. The assumption is made here that a gall midge larvais present and will be recognized as such, usually by the factthat it possesses an anchor-process or breast-bone (PI. i).This is followed by brief sections covering damage, descriptionand distribution. Experience has shown that it is frequently

P reface . 9more or less useless, and usually most unwise, to attempt toidentify a species from keys with any certainty unless biologicaldata are available in addition. Particularly would this be trueif non-specialists attempted specific determination on caughtspecimens. To emphasize and avoid this the briefest possibledescription is given, but reference is always made to theoriginal description.Keys for the separation of the genera, and sometimes alsothe species, are available in the works of Kieffer (io8). Felt(69) and Riibsaamen & Hedicke (178) for students who wishto specialize on the gall midges.The life history is next considered in detail as it occurs inthe different countries in which it has been studied. Thenfollow sections on food plants and natural enemies. Control isnext dealt with fully. Then, if the species is represented in mycollection of permanent slides, the material present and itssource are given.Lastly, there is a list of references, the first always being thatof the original description, followed if necessary by those of thespecies' synonyms, and then in chronological order the mostimportant papers dealing with the biology or control. For somespecies additional sections dealing with synonyms, fluctuationsin numbers, and inquilines, or subdivisions of the previouslymentioned sections are inserted. For many species, however,the available information is not sufficient for any side-headingsat all. The text is fully annotated in order that anyone mayreadily pursue any matter further.Each volume will be concluded by a list of references, anindex to the generic, specific and popular names of the gallmidges mentioned, a plant index and a general index.It will quickly become obvious to any reader that the lifehistory of many of the species still remains to be worked out indetail. In addition, it will be seen that the gall midges as a groupprovide ample material for the study and elucidation of suchproblems of general interest as periodicity in outbreaks ofinsect pests, host plant range of phytophagous insects, rangeof prey ôf the 'zoophagous species, and soil conditions thatenable the larvae of certain pests to remain several winters in

10 Gall Midges of Economic Importancethe soil before pupation and emergence of the perfect insect,and so on. • iIn fact, throughout the period I have been studying thefamily I have been attempting to follow the adage so succinctlypropounded by G. F. Ferris (73),: "The systematist may andshould employ any means that are available in order to arriveat a knowledge of the biological facts, whether those meansmay be found in morphological, anatomical, physiological,experimental, genetical or even chemical studies." Likewisethe worker in applied entomology should use all the weaponsof knowledge that are available in order to arrive at a trueunderstanding of the insect he is called upon to control andprevent reaching epidemic numbers.When starting an investigation it has always .been foundadvisable to wait if possible until the larvae are nearly fullgrownbefore collecting them. Then the part of the plant onwhich they are, or with small plants the whole plant, shouldbe collected and placed in a simple breedmg cage. This consistsessentially of a flower-pot containing soil surmounted bya layer of fibre (coco-nut, peat, or bulb fibre without carbon),and either a lamp glass covered by musUn sewn on to an ironring or a muslin bag supported on bamboos or wire. Thepots are placed in saucers.Thus if a midge is infesting grass-heads, fifty or a hundredof the latter may be placed in a lamp glass containing fibreand standing on the soil in a 5-inch flower-pot. For more bulkymaterial, such as twigs or leaves, it is advisable for the flowerpotto be of the 8-, 10- or 12-inch size, the layer of fibre to coverthe soil to a depth of 2 to 3 inches and the whole to be enclosedby a muslin cage. The height of this depends on the materialto be enclosed. The base of the cage can easily be attached tothe top of the pot by tying with string. Thus if the larvae leavethe galls to pupate, they can descend to the fibre; if they dcfnot, they can remain in the plant material.It has been found that damp fibre provides an excellentmedium for the survival of the larvae until pupation of thosespecies that normally descend to the soil at this stage in theirlife history. Soil, on the other hand, is most unsatisfactory

P r e'f a c eowing to its liability to cake or dry out so much more rapidlywhen kept in such small volumes. It should be realized thatmany species pass the winter as larvae in the soil and do notpupate until shortly before emergence.The cages described have been kept in an outdoor insectarythat is unheated and of which the roof is so designed as to allowthe sun to reach the pots on the shelves only for a few hourseach day. Owing to this the difficulty of moisture condensingon the inside of the lamp glasses is reduced to a minimum.On the emergence of the midges they are removed one byone with an alcohol-wetted paint"brush if desired as specimens,or in glass tubes if required alive. With a little practice onesoon becomes accustomed to removing midges alive withoutallowing any to escape. Naturally it is much more easy toremove midges when there are only a few newly-emerged.They assemble on the muslin at the top of the cage. Thisrequires frequent inspection of the cages so as not to arrive atthe cage for the first time well after emergence has started.Another advantage to b'e gained by frequent inspection's thatone can remove the midges before they have mated, thusobtaining material for cross-mating experiments, etc. Whenthe midges have been tubed alive, these tubes are either stoodin sand or plugged with cotton wool. Care must be taken notto allow direct sunlight to fall on them.When sufficient live midges have been collected they maybe allowed to mate in the tubes or be inserted direct into amuslin-covered flower-pot containing their host plant, which,of course, is grown in an ordinary potting mixture of soil, fibreand sand. It is advisable to roll up the base of the muslin cageon to the top of the flower-pot so that the top of the cage isabout 2 inches above the top of the enclosed plant. The cageis raised by unrolling the muslin as the plant grows in height.The midges are best inserted through a hole cut in the sideof the muslin cage, the hole being afterwards plugged withcotton wool.In the spring care must 'be taken to prevent sparrows, etc.,removing the wool plugs for nesting purposes, and again in theautumn a sharp watch must be kept for holes in the muslinII

12 Gall Midges of Eco'nomic Importancecages, made by wasps eating their way in. With a little practicethe movements of the midges pside can. be observed, as wellas their mating and egg-laying. If the larvae are expected todescend to the soil for pupation, it is as well, though notessential, to provide a layer of fibre on the top of the soilmixture. Sand, however, should on no account be used asa substitute for the fibre. When the next generation of adultmidges appears they can be removed by inserting test-tubesthrough the holes in the muslin.By using the above simple methods it has been possible torear successfully large numbers of midges. When only rearingto obtain material for distribution or fluctuation studies andso on in which only dead material is needed,! have been ableto handle up to 5,000 midges in a single day and go on dealingwith 3,000 a day for 8 weeks or so.If, however, the midges are required for breeding experimentsit has usually only been possible to set up experimentsneeding 50 or so live midges a day and the rest of the day'semergence has to be extracted from the cages and used aspickled material. Midges live as adults only for a few daysand consequently it is always advisable to use midges forbiological experiments on the day they emerge. Since midgesemerge at fixed periods in the day and different species havedifferent times of emergence in the 24 hours one can deal comfortablywith several species the same day.Further, after a little experience one can avoid investigatingspecies that all have the main emergence at the same time ofthe year, and so one can stagger the rush periods. The realkey to success in breeding gall midges, however, as is true withall animals, is to have patience and enough time to makecountless observations and minor adjustments. Insects will notbreed normally or well'under artificial conditions if just putin cages with plenty of food and then left to themselves withonly irregular and infrequent inspection.The larvae, pupae and midges may be preserved in a satisfactorycondition for years by placing them direct (but themidges not before the wings have hardened) into tubes containinga mixture of 80 per cent, alcohol and a small quantity

Preface 13(at the rate of a drop or two per small tube) of pure glycerine.The strength of the alcohol allows it to decrease in time to70 per cent., while if it evaporates entirely the glycerineremaining at the bottom of the tube will preserve the materialuntil the next inspection. In such instances the material canbe made available for study by simply refilling the tube withfresh alcohol. It is necessary to stress the fact that each tubeshould be fully labelled, since so much valuable material isrendered useless by omitting to carry out this elementary rule.Material kept in this mixture has been found in perfectcondition for taxonomic study after 15 to 20 years.When time allows, the material should be mounted as microscopeslides. Gall midges mounted on cards or pin-pointsbecome worthless very quickly. Here a word of warning isessential. If one attempts to make slides of midges too soon ,after their emergence the colour pigments are apt to diffuseout of the body and. stain the preparation. It is best to allowthem to remain in the alcohol for 3 to 4 weeks after emergence.Similarly, if the alcohol is too strong bleaching will take placevery quickly. The best strength is about 75 per cent.When mounting the midges, they can be removed from thealcohol-glycerine mixture and placed in 95 per cent, alcoholand allowed to remain there for about i hour, depending ontheir size and freshness. They are next placed in carbol-turps(carbolic acid crystals 40 per cent., rectified spirits of turpentine60 per cent.) for not more than a minute or so and then transferredto a slide. Then, using fine needles, the head should beseparated from the body, and the genitalia of the males also.The carbol-turps must then be drained off with filter pkperand Canada balsam substituted. It is important to drain offthe carbol-turps since it does not mix with Canada balsam.The head and genitalia are separated from the rest of themidge in order that they may be mounted flat.The same process can be followed for mounting whole pupae,empty pupal cases, or even whole larvae. For larvae, however,it is very important to mount larval skins. This can be done bypricking the larvae with a fine needle and boiling in potashuntil the body contents have softened. Then the latter are

• Preface 15written on the labels, whicK are later painted red. For paratypesgreen is used, while for cotypes green and red are used.Homotypes are labelled as such. Each slide is represented ina card index by a card bearing the data on the slide labels andany extra information. These cards are kept in numericalorder. In addition there are indexes of species and genera, thecards of which indicate respectively which numbers in thecollection represent the species and which species representthe genus.When describing species or referring to a particular-specimen,it has been my practice to refer to the slide number in theBarnes collection.The amount of time taken in following this routine whenmaking slides is amply repaid when examining slides madesome years previously.Various workers, namely Dr. Margot E. Metcalfe, MissJoan A. T. Anderson and Dr. D. P. Jones, have deposited inthis collection the original material of the species they havedescribed. Numerous other entomologists throughout the worldhave been kind enough to send material for the collection.I have also been permitted to retain in the collection materialsent to me for identification by the Imperial Institute ofEntomology. I feel that this is an appropriate place to expressmy thanks to all those who have made it possible to build upthis representative collection.My thanks are due to the University of Leeds and theYorkshire County Council for permission to reproduce thecoloured figures from the late Mr. T. H. Taylor's bulletin onthe.swede midge; to Dr. Kenneth M. Smith, F.R.S., and theCambridge University Press for Plate i and Fig 2, Plate 6(from K. M. Smith's Textbook of Agricultural Entomology); tothe Royal Horticultural Society and to C. A. W. Duffield, Esq.,who took the original photograph, for Fig. i, Plate 6 (from theJournal of the Royal Horticultural Society); to the Ministry ofAgriculture for Plate lo (from the Journal of the Ministry ofAgriculture); to *Dr. E. R. Sp'eyer, Entomologist to the CheshuntExperipjental Station, for the photographs of Mycophilaspeyeri and Pezomyia vanderwulpi; and to Mr. V. Stansfield

16 Gall Midges of Economic Importancefor taking the photographs of the damage done to parsnipleaves by Macrolabis corrugans. I am indebted also to Mr. D. C.Thomas, of Seale Hayne Agricultural College, for permissionto use the typescript account of his current work on the swedemidge.Finally, I would like to take ithis opportunity of expressingmy gratitude to all my entomological friends and colleagueswho are too numerous to name individually but who have beenan unending source of encouragement and help in the preparationof this work. It is my hope that in return it may beof some slight assistance to them.I am indebted to Mr. C. T. Gimingham for writing theForeword.H. F. BARNES.ROTHAMSTED EXPERIMENTAL STATION.January, 1945.

Contents of Vol. IPageForeword by C. T. Gimingham, O.B.E., B.Sc, F.R.I.C. . . . 5Preface 7Contents 17List of Illustrations .19Introductory 21Root and Vegetable Crops attacked by Gall Midges . . . 22Asparagus (Asparagus officinalis L.) 24Bitter Gourd (Momord/ca spp.) 25Broad Bean (Wc/a faba L.) 26Carrot (Caucus carota L.) 26Cassava (Manihot utilissima Pohl and M. aipi Pohl) . . . 30Coednia indica (Naud.) Wight & Arn 33Crataeva religiosa Forst . . . . . . . . . 34Cruciferous Crops (Cabbages, Turnip, Swede, Rape, Radish, etc.) 34Egg-plant or Aubergine (So/anum mefongena L.) . .57Leek (Allium porrum L.) . . . 57Lentil (Lens esculents Moench) 58Lettuce (Lactuca satiVa L.) 59Mushroom (Psa//ota (Agaricus) eampestrls Qu6l. or P. (A.) arvensis Quî) 60Parsnip (Past/naca satlva L.) 64Pea (Pi'sum satiVum L. and P. arvense L.) 68Potato (So/anum tuberosum L.) 76Rhubarb (Rheum rhaponticum L.) • 77Squash (Cucurbita pepo L. var.) 77Sweet Potato (Ipomoea batatas Lam.) 77Tomato (So/anum /ycopers/cum L.) 78Vegetable Marrow (Cucurbita pepo L. var.) 8017 B

18 Gall Midges of Economic ImportancePageYam (O/oscorea spp.) 82References . i 83Index of Generic, Specific and Popular Names of Gall Midgesmentioned . . \ 94Plant Index 98General Index 101

List of IllustrationsPlate I.Facingpage"Anchor Processes" or "Breastbones" of various Cecidomyldlarvae. Greatly enlarged. I. Cam()y/omyza ormerodi; 2. Dasyneuraleguminicola; 3. Sitodiplosis mosellana; 4. Clinodiplosisp/si'co/a; 5. Contarinia pisi; 6. Dasyneura trifolii; 7. Mayetioladestructor; 8. Contar/n/a tritici. (After K. M. Smith) . . 8Plate 2. Fig. I. Male Swede Midge, Contarinia nasturtii. Magnified.(After T. H. Taylor)Fig. 2. Female Sv/ede Midge, Contarinia nasturtii. Magnified.(After T. H. Taylor) 35Plate 3. Fig. I. Swêde Midges on a young Swede Leaf. About natural size.(After T. H. Taylor)Fig. 2. Eggs of Swede Midge on upper surface of Stalk of youngSwede Leaf. Natural size. (After T. H. Taylor)Fig. 3. Eggs of Swede Midge on upper surface of Leaf-stalk.Magnified 30 times. (After T. H. Taylor) . . 39Plate 4. Larvae of Swede Midge on upper surface of Stalk ofyoung Swede Leaf. Magnified 4 times. (After T. H.Taylor) 41Plate S. Fig. I. Pupal Cocoon of Swede Midge. The inset shows naturalsize of cocoons. (After T. H. Taylor)Fig. 2. Pupa of Swede Midge. Magnified 45 times. (After T. H.Taylor) 43Plate 6. Fig. I. Brassica siliqua infested with Larvae of the Brassica PodMidge, Dasyneura brassicae. (Photograph by C. A. W.Duffield)Fig. 2. Larvae of the Pea Midge, Contar/n/a p/si, in pea pod (x U).(After K. M. Smith) . . . . . . 5 2Plate 7. Fig. j. 1. Male Mycophila speyeri (x 50)Fig. 2. Female Mycophila speyeri (x 50) . . . . . 61(Photographs by E. R. Speyer)Plate 8. Fig.I. 1. Female Pezomyia vanderwulpi (x SO)Fig. 2. Egg Mycophila speyeri (x 70)Fig. 3. Larva Mycophila speyeri (X 72) 62(Photographs by E. R. Speyer)Plate 9. Figs. I and 2. Leaves of Parsnip infested by Larvae ofMacro/ab/scorrugans. (Photographs by V. Stansfield) . 65Plate 10. Fig. I. Pea Flowers infested by Larvae of the Pea Midge, Contar/n/apisiFig. 2. Pea Shoot infested by Larvae of the Pea Midge, Contariniapisi 6819

IntroductoryI n Vol. I an attempt has been made to deal withall the gall midges that have been recorded as pests of rootand vegetable crops. It will be readily recognized that thegall midges that attack the more common vegetables havereceived more attention from economic entomologists thanthose that live on plants only used locally as vegetables. Itis for this reason that some of the gall midges that infest therarer vegetables may have been overlooked. In some instancesmidges that have not actually been reported as pests have beendealt with, since they may easily become pests in the future.The damage done may involve several parts of the plant,e.g. the pea midge {Contarinia pisi Winnertz) and the swedemidge (C. nasturtii Kieffer). On the other hand, it maybe restricted to the seeds, e.g. the carrot midge {Kiefferiapimpinellae F. Loew), and the brassica pod midge {Dasyneurabrassicae Winnertz). Others prevent the flowers opening andso indirectly forestall any chance of fruit, e.g. the tomato flowermidge (C lycopersici Felt). In still other instances only theleaves and terminal shoots are attacked, e.g. the cassava midgesand the vegetable marrow midge [Jaapiella bryoniae Bouche).With at least two species the root development is materiallylessened as a consequence of midge attack, i.e. in swedes bythe swede midge (C. nasturtii Kieffer) and in parsnips byMacrolabis corrugans F. Loew. Thus the damage to the crop maybe direct or more commonly indirect.The alphabetical list of the crops dealt with in this volumeis given in order to provide a clue to the identification of themidges. It indicates the gall midges that may be involved andthe part of the plant they attack. Subsequently the midgesare considered under the vegetables they attack.21

ROOT AND VEGETABLE CROPS ATTACKEDBY GALL MIDGESAepim, Sweet Cassava or Mandioc {Manihot aipi Pohl)Jatrophobia brasiliensis Riibsaamen Leaves, p. 32Asparagus {Asparagus officinalis L.)Contarinia florum Riibsaamen Flowers, p. 24Unidentified gall midges Leaves, branches, stems, p. 24Aubergine, see Egg-plantBitter Gourd {Momordica spp.)Dasyneura beccariella Del Guercio Flowers, p. 25Lasioptera falcata Felt Stems, p. 25Unidentified gall midge Stems, p. 25Broad Bean {Vicia faba L.)Contarinia pisi (Winnertz) Pods, p. 68Prqfeltiella sp. Stalks, p. 26Cabbage {Brassica oleracea L.), see Cruciferous cropsCarrot {Daunts carota L.)Kiefferia pimpinellae (F. Loew) Fruit, p. 27Lasioptera carophila F. Loew Umbellules, p. 26Cassava or Mandioc, Bitter {Manihot utilissima Pohl)Cecidomyia manihot Felt Leaves, p. 31Jatrophobia brasiliensis Riibsaamen Leaves, p. 32 •Lasioptetyx manihot Felt Stems, p. 30Schizomyia manihoti Tavares Leaves, p. 31Cassava or Mandioc, Sweet, see AepimCauliflower {Brassica oleracea L. var.), see Cruciferous cropsCoccinia indica (Wight & Am.)Neolasioptera cephalandrae Mani Stems, p. 33Colza {Brassica napus L.), see Cruciferous cropsCrataeva religiosa Forst.Cecidomyiella crataevae Mani Leaf buds, leaflets, p. 34J/eolasioptera crataevae Mani Flowers, p. 34Cruciferous crops (Cabbages, Turnip, Swede, Rape, Radish, etc.)Contarinia nasturtii (Kieffer) Flowers, leaves, shoots, p. 35Dasyneura brassicae (Winnertz) Siliquas, p. 51Gephyraulus raphanistri (Kieffer) Flowers, p. 55' 22

Gall Midges of Economic Ir^portance 23Egg-plant {Solanum melongena L.)Asphondylia sp. Unknown, p. 57? Contarinia lycopersici Felt Flowers, p. 57Unidentified gall midge (in Eritrea) Shoots p. (57Unidentified gall midge (in B.W.I.) Flowers p. 57Leek {Allium porrum L.)Clinodiplosis sp. Seed heads, p. 57Lentil {Lens esculenta Moench)Asphondylia ervi Riibsaamen Pods, p. 59Dasyneura viciae (KiefFer) Leaves, p. 59Unidentified gall midge p. 58Lettuce {Lactuca sativa L.)Clinodiplosis sp. Seed heads, p. 59Mushroom {Psalliota campestris Quel, or P. arvensis Qu^l.)Lestremia cinerea Macquart "Stalks, p. 64Miastor sp. Mycelium, gills and cap, p. 6gMycophila barnesi Edwards Mycelium, p. GjMycophila fungicola Felt Young mushrooms^ p. 62Mycophila speyeri (Barnes) Mycelium, p. 6(Parsnip {Pastinaca sativa L.)Contarinia pastinacae (Riibsaamen) Fruit, p. 5^.Kiefferia pimpinellae (F. Loew) Fruit, p. 27Macrolabis corrugans (F. Loew) Fruit, p. 64Pea {Pisum sativum L. and P. arvense L.)Contarinia pisi (Winnertz) Flowers, ^shoots, pods, p. 68Potato {Solanum tuberosum L.)Asphondylia trabuti Marchal Fruit, p. 76Radish {Raphanus sativus L.), see Cruciferous cropsRape {Brassica napus L.), see Cruciferous cropsRhubarb {Rheum rhaponticum L.)? Profeltiella sp. Stalks, p. 77Sepim {Manihot palmata Miill.-Arg.)Jatrophobia brasiliensis Riibsaamen Leaves, p. 03Squash {Cucurbita pepo L. var.)Cecidomyia cucurbitae (Felt) Surface of fruit, p. 77Swede {Brassica rutabaga), see Cruciferous cropsSweet Potato {Ipomoea batatas Lam.)Schizomyia ipomoeae Felt Flower buds, p. 74Tomato {Solanum lycopersicum L.)Contarinia lycopersici Felt Flowers, p. 78Turnip {Brassica campestris L.), see Cruciferous crojjgVegetable Marrow {Cucurbita pepo L. var.)Jaapiella bryoniae (Bouche) Terrriinal leave^^ p. 80Yam {Dioscorea spp.)Unidentified gall midges Leaves and stem^ p. 82

ASPARAGUS {Asparagus officinalis L.)No report has been published in economic Uterature of gallmidges being injurious to asparagus (Asparagus officinalis L.),although at least two species are known to occur on this plantin Europe.The larvae of Contarinia florum Riibsaamen live gregariouslyin the blossoms, which remain closed and swollen. Pupationtakes place in the soil. Riibsaamen (176) stated that thelarvae were white and described both sexes from Germanmaterial; but Ross & Hedicke (158) referring to this speciesstated that the larvae were orange in colour. Dittrich &Schmidt (47) also referred to orange-coloured larvae in unopenedblossom.There are also records of unidentified gall midge larvaeliving in little swellings on the stem in Central Europe (104)and in similar swellings localized at the axils of young branches •in Italy (218).Two other species of gall midges have been described fromother species of Asparagus; viz. Dasyneura asparagi (Tavares) inPortugal on A. aphyllus L. and D. turionum (Kieffer & Trotter)in Algeria, Italy and France on A. acutifolius L.Tavares (194) described the larva and both sexes of D.asparagi. The larvae are white, gregarious and pupate in thesoil. They live between the leaf-spines, which are clustered,twisted, discoloured and swollen at the bases, at the extremitiesof young branches (195).Kieffer {in 216) described both sexes and the larvae ofD. turionum. Its larvae are flesh coloured and live eithersingly or several together,, probably pupating in the soil.24

Gall Midges of Economic Importance 25Their galls are similar to those of D. asparagi, being clusters ofleaves swollen at their bases, discoloured or reddish, at the endsof young branches that are swollen and twisted (86, 132, 215,217)-Other unidentiiied gall midge larvae living on the leaves ofA. albus L. and A. horridus L. have been reported from Algeriaand Tunisia. For references Houard (87) may be consulted.BITTER GOURD {Momordka spp.)Two species of gall midges have been reported on Momordicacharantia L., namely Lasioptera falcata Felt and an unidentifiedspecies.L. falcata was originally described (66) on a single femalereared from a stem gall on an unidentified wild cucurbit in thePhilippines. Felt later referred specimens of a midge bred onM. charantia in South India to this species. Ramakrishna (154)mentioned this midge as the bitter gourd vine gall fly andfigured the gall and adult insect. The galls, which are longand tubular, are to be found on the distal young shoots. Theyaffect the growth of the plant. Mani.(i27) stated that it wasvery common at Coimbatore and had been found in variousother localities in South India. No description of the malehas apparently yet been made, neither has an account beenpublished of its biology.Docters van Leeuwen-Reijnvaan (49) described from Javaand Celebes a gall on the stem of M. charantia. This is up to100 mm. in length and contains numerous larval cavities, eachof which contains an orange Cecidomyid larva. It resemblesthe stem gall which was figured by Docters van Leeuwen-Reijnvaan (48) and is caused by Prolasioptera javanica (Kieffer& Docters Reijnvaan) (no). The larvae of this last-namedmidge cause swollen flower and stem galls on Melothria perpusillaCogn. in Java and Celebes. For additional referencesto this midge see Houard (87).Another species, Dasyneura beccariella Del Guercio, has beenfound on Momordica pterocarpa Hochst. (pteromorfa) in Eritrea.The larvae, female and gall were described (46). The infested

26 Gall Midges of Economic importanceflowers remain closed and in a bunch. The larvae are yellowishorangeand gregarious. Del Guercio gave figures of the larvaand gall. He also mentioned an unidentified Chalcid parasite.I *BROAD BEAti{Vicia faba h.)The pea midge {Contarinia pisi Winnertz) has been recordedon this plant (see p. 72). In addition, Bagnall & Harrison (7)recorded finding red larvae in galls similar to those ofAsphondylia lathyri Riibsaamen (173) in the pods of cultivatedbeans in Durham.A Profeltiella species also lives on broad bean. In August,1941, J. H. Stapley found larvae of this genus inside the stalksof broad bean in the Cambridge province, England. Femalemidges were reared from the larvae by the author. Both larvaeand females are in the Barnes collection. The larvae of thetype species, P. ranunculi KiefTer (105), of this genus are statedto feed with the larvae of another gall midge^ Geodiplosisranunculi KieflFer (105), on the roots of Ranunculus acer inGermany and England (6). Another species, which may bewrongly placed generically, is P. orientalis Felt (64), which wasreared in association with the gall midge Kamptodiplosis reductaFelt (64) from leaf galls on Siphonodon celastrineus in thePhilippine Islands. Two other species, P. dizygomyzac Barnes andP. vespicoloris Barnes, will be considered in a later Volume.The remaining species, of the genus is P. soya (Monzen), whoselarvae live in the petioles and stems of the soy bean in Japanand are considered a pest. This species will be dealt with inVol. II.CARROT {Daucus carota L.)Two gall midges have been recorded as attacking this plant,viz. Lasioptera carophila F. Loew, whose larvae live in galls onthe umbellules, and Kiefferiapimpinellae (F. Loew), whose larvaelive in the swollen fruit. L. carophila will be dealt with in alater volume as a caraway pest, since it is on this plant thatit has been recorded as a -pest.

Gall Midges of Economic Importance 27Kiefferia pimpinellae (F. Loew, 1874), the Carrot GaU Midge,*syn. pericarpiicola Bremi, 1847 (laiva and gall only), dauci' Bremi, 1847(larva and gall only), umbellatarum F. loew, 1877.DIAGNOSTIC CHARACTER. Swollen fruit, standing out wellabove the inflorescence, of Carrot and Parsnip.DAMAGE. The fruits become reddish or violet in colour andhave a tendency to lose their normal shape and spines. As aresult of an infestation no seed is formed.There are only two records of this midge being of economicimportance. Noel (146) listed it as a pest of parsnip in France,while Rostrup & Thomsen (162, 163), referring to it asAsphondylia umbellatarum, stated that, although it had often beenfound on wild carrot, it had only once been found as a pestof cultivated carrots. This was in Denmark in 1924, whenpractically all a seed crop was rendered useless.DESCRIPTION. F. Loew (123) described this species, whichwas originally in the genus Asphondylia and later transferred byRiibsaamen (167) to the genus Schizomyia. Mik (140) placedit in his newly-erected genus Kiefferia as the generic type. Itmay be distinguished from Asphondylia and Schizomyia speciesby the ovipositor having two lamellae at its extremity insteadof being entirely needle-shaped. The larvae is yellowishorangeand has a pointed anchor process. Its anal segmentdoes not have the two curved processes typical of Schizomyiaspecies.^SYNONYMY. Bremi (32) suggested the name Cecidomyiapericarpiicola for this species, knowing and figuring only thelarva and gall, but in the legend he refers to it as C. dauci.F. Loew (124) changed the name pimpinellae to umbellatarumbecause he had found it on so many species of Umbellifers.Riibsaamen (169) stated that Bremi and H. Loew had referredto the galls of the same midge under the names C. pericarpiicola,C. dauci, C. pimpinellae and C thysselini.DISTRIBUTION. Generally throughout Europe, includingHolland (136), Germany (loi, 123), Italy (131), France (150),Switzerland (H.F.B. coll.), Hungary (193), Portugal (195),Sicily (190), Finland (122), Denmark (162) and Russia (169).* Also Mohrengallmucke, Gulerods-galmyggen.

28 Gall Midges of Economic ImportanceIt has also been recorded in England, by Bagnall & Harrison (3)among others, on Angelica sylvestris, Pimpinellae saxifraga andDaucus carota and in Scotland by Binnie (27) on Angelicasylvestris. It is probably widespread in the British Isles on wildUmbellifers.LIFE HISTORY. Exceedingly little has been reported in theliterature concerning the biology of this species. The yellowishlarvae live singly in the swollen fruit. The galls are to be foundin the middle and late summer and in August the larvae arefull grown and descend to the soil where they spend the winteras larvae in cocoons. F. Loew (123) reporting that he still foundthem in the larval stage in March. The midges start appearingat the end of May. There is probably only one generation ayear.FOOD PLANTS. Besides being found on Caraway (Carum carviLi.), Carrot {D. carota L.) and Parsnip {Pastinaca sativa L.)this midge has been recorded from many Umbelliferous plants.These include wild Angelica [Angelica sylvestris L.), Chervil{Anthriscus silvestris Hoffm.), Burnet Saxifrage {P. saxifraga h.),the plant from which it was described by F. Loew, Fennel[Foeniculum officinale All.), Upright Hedge-Parsley [Torilisanthriscus Gmelin), Sickle-leaved Buplever {Bupleiirum falcatumL.), B. ranunculoides L., B. longifolium L., Falcaria rivini Host,Pimpinella magna L., P. tragium Vill., Annual Meadow Saxifrage[Sesili annuum L.), Foeniculum piperitum Sweet, Silaus pratensisBesser, Peucedanum oreoselinum Moehch, P. palustre Moench,Cow-parsnip or Hogweed [Heracleum sphondylium L,), Laserpitiumpruthenicum L. and Daucus carota var. prostratus Rouy &Camus.It has also been doubtfully identified as occurring on Parsley[Petroselinum sativum HofFm.) and Pkysospermum aquilegifoliumKoch.The authorities for these records can be ascertained onreferences to Houard (85).NATURAL ENEMIES. Kieffer (log) described Synopeas daucicolaas a larval parasite.INQUILINES. Four gall midges have been reported as livingin the galls of Kiefferia pimpinellae, namely Trotteria umbelliferarum

Gall Midges of Economic Importance 29(KiefFer), Amerhapha gracilis Riibsaamen, Trotteria inquilinaRiibsaamen and Contarinia inquilina Rubsaamen.KiefFer (104) first mentioned T. umbelliferarum in a footnotedescribing the male very briefly as a species of the genusChoristoneura Riibs. Later in the same work he replaced thisgeneric name, because it had already been used to denote aTortricid genus, by the new name Trotteria. KiefFer thenrecorded it in galls on Daucus carota, Pimpinella magna andPeucedanum oreoselinum in Lorraine. Its larvae may be predaceouson the larvae of Kiefferia pimpinellae. Later (106) KieflFer gavea more extended description of the male and female, statingalso that the larvae are rose-red in colour and live in the gallsof K. pimpinellae on different species of Umbelliferous plants atBitche. Bagnall & Harrison (3) reported it on Anthriscus inDurham, England.A second species of Trotteria, T. inquilina Rubsaamen, hasalso been recorded from the galls of K. pimpinellae on Sesili(Libanotis) montana in Germany. Rubsaamen (177) describedboth sexes.Amerhapha gracilis, whose larvae are orange-red to red, hasbeen recorded by Bagnall & Harrison (7) from Shropshire,England. These workers state that the larvae may be found .fully fed in mid-October after those of Kiefferia pimpinellae haveleft the galls. Only the male was described by Rubsaamen(173), who mentioned that the orange-red larvae had beenfound in Germany living in the galls of K. pimpinellae. At thesame time he erected the genus Amerhapha with gracilis as thetype species.The larvae of Contarinia inquilina, which are orange-yellow,were reported by Riibsaamen (176) as living in the galls ofK. pimpinellae in Germany.CONTROL. No control for the carrot gall midge has yet beensuggested: its occurrence as a pest is too rare an event.MATERIAL. Larva, Switzerland (from Daucus carota).

30 GallMidges of Economic ImportanceMOST IMPORTANT REFERENCES*LoEw, F. 1874. Verh. zool.-hot. Ges. Wien, 24, 326-8 {Asphondylia)RosTRUP, S. & THOMSEN, M. 1928. Vort landrugs skadedgyr, p. 251 {A.umbellataruni) ,RosTRUP, S. & THOMSEN, M. 1931.J Die tierischm schddlinge des ackerbaues,p. 272 {A. umbellatarum).CASSAVA {Manihot utilissima Pohl and M. aipi Pohl)Cassava is extensively cultivated in the tropics for the largetuberous roots, which contain a large amount of starch. Theroots, when ground, form mandioc or cassava meal, sometimescalled Brazilian arrowroot, f By a special process tapioca isprepared from the roots. Cassava root, flour (mandioca flour)and starch are used in the English textile industry. The driedroot is also a source of industrial alcohol. Even the poisonousjuice, when extracted and evaporated, thus having beenrendered harmless, is used as an antiseptic, cassareep, in preservingmeat.Some authorities consider that Bitter Cassava is M. utilissimaPohl and that Sweet Cassava is M. aipi Pohl {M. palmataMiiU.-Arg.). In Sweet Cassava most of the hydrocyanic acidis confined to the rind of the root, whereas in Bitter Cassavait is fairly evenly distributed in the core and rind of the root.This character, however, is not constant in different locaUties.For discussion of these points reference should be made toSampson (180).Four species of gall midges have been described from leafgalls of cassava, viz. Cecidomyia manihot Felt, Lasiopteryx manihotFelt, Jatrophobia brasiliensis Riibsaamen and Schizomyia manihotiTavares.L. manihot was described (61) as a small yellowish midgereared from M. utilissima by W. H. Patterson in St. Vincent,B.W.I., and erroneously stated by Felt (65) to have been reared* In this section the first quoted reference is, throughout the work, that of theoriginal description. Unless there are references to the original descriptions ofsynonyms, the remaining references are the most important references to thebiology and control of the midge in question.•f The true arrowroot of commerce comes from the rhizome of Maranta arundinacea."

Gail Midges of Economic Importance 31from leaf galls. Patterson (149), however, states that thisspecies was obtained from decaying material in cassava stemsattacked by weevils. Urich (220) wrote that the red growths orgalls often seen in Trinidad on the leaves of cassava were causedby a small yellowish midge, Lasiopteryx sp. It is most probablethat Urich should have referred to these galls as the work ofCecidomyia manihot Felt or Jatrophobia brasiliensis Riibs. With thispossible exception no additional record of L. manihot has beenmade in the literature.Schizomyia manihoti was described and figured in detail byTavares (203). The galls of this species are globular, aboutthe size of a millet grain, glabrous and reddish; they areinserted on the side of the mid-vein of the leaves of M. utilissima.Most often the galls are agglomerated forming masses,7-40 mm. X 10-15 mm., which occupy nearly all the leafsvirface. NumeroMS larval cavities, each contftiniiig a red larva,3 mm. long, are to be found in such a mass. Pupation occursin the galls. So far this species has only been recorded fromBrazil (by Tavares) and apparently has not appeared againin the literature.Cecidomyia {Itonida) manihot was described by Felt (57) as asmall yellowish-brown species, only 1-1-25 "^- long. Bothsexes were described. He stated it was reared by W. H.Patterson, St. Vincent, B.W.I., from leaf galls on M. utilissima.The same authority (68) stated that he had also receivedspecimens from F. W. Urich in Trinidad during March, 1915.Urich himself mentioned (220) an unidentified Lasiopteryxspecies (see previous paragraph but one). Several workers havesubsequently mentioned C. manihot Felt. For example. Watts(223) stated that it was extensively parasitized, and that twospecies of minute Hymenoptera had been reared in St. Vincent,while Harland (81) recorded Tetrastichus fasciatus Ashmead asa larval parasite. Girault (77) gave a description of this insect.Donovan (51) said it was present throughout Grenada. BothWatts and Harland stated that the midge did very little harm,but Bequaert (26) stated that it was probably the mostdangerous pest of cassava in South America, His figures of thegalls remind one forcibly of those of Jatrophobia brasiliensis

32 Gall Midges of Economic importanceRiibs., and although he quotes Felt's description of C. manihot,he probably was dealing with the next species, J. brasiliensis.J. brasiliensis was originally described from Brazil on thelarva, pupa and gall only by'Rubsaamen (171) as Clinodiplosisbrasiliensis on Manihot utilissima and M. dichotoma Ule. Later,he (174) erected the genus ^latrophia [sic), better written asJatrophobia after the old generic name of cassava, and describedboth sexes of the adults under the name /. brasiliensis. Tavares(199) received galls from Brazil on M. utilissima and M. aipi,figuring and describing them in detail. Apparently unawareof this later description by Rtibsaamen, he described the femalemidge, placing it in the genus Eudiplosis as E. brasiliensis Riibs.Still later, Tavares (200), discovering that the name Eudiplosishad already been used by Kieffer (100), changed the name ofhis genus Eudiplosis to Autodiplosis. The result of these frequentname changes is ihsX brasiliensis Riibs. is referred to in literatureunder the four generic names. It appears most reasonable torefer to it as J. brasiliensis Riibs.This species forms nail-shaped galls on the upper surface ofthe leaves of M. dichotoma Ûle, M. aipi Pohl [M. palmataMiill.-Arg.) and M. utilissima Pohl. The galls * are 11-15 mm.tall and 3-5 mm. wide, glabrous, yellowish or pale green andsometimes reddish. They open on the lower surface of theleaves. A single larva, when full grown 1-2-2 mm. in size,lives in each. The entire life-cycle is spent in the galls which aremost abundant near the mid-vein. Bondar (28, 29) gives briefdescriptions of the life-cycle and states that usually little harmresults, but that occasionally the galls are numerous enough tocause the leaves to become deformed. In such instances theleaves should be destroyed, as the growth of the plant is retarded.This species is definitely known from Brazil, BritishGuiana, Trinidad and Tobago (36) and also St. Vincent (37).Myers (142) studied two of its parasites; the Ghalcidoids,Aprostocetus fidius Gir., which is a brown species, and an unidentifieddark metallic blue species of the same genus. Callan(37) stated that the

Gall Midges of Economic Importance 33iVincent and the Eulophids Aprostocetus fidius Gir. and Aprostocetus -sp. in Trinidad were on record, while he himself had bredTetrastichus fasciatus in Trinidad and British Guian^.Specimens in the author's collection are as follows: males,females and pupae, Trinidad and British Guiana; larvae,British Guiana. These all agree with Riibsaamen's descriptionof^. brasiliemis.There is some doubt about the differentiation of these twolast-named species, C. manihot Felt and J. brasiliensis Riibs.Rubsaamen (174) said that they did not appear to be identical,and Felt (69) also accepted J. brasiliensis Riibs. and said thatthe two species could not be identical if Riibsaamen's genericplacing was correct. Houard (88), however, gives Felt'sreferences to C. manihot bracketed amongst the references toJ. brasiliensis and also includes in the list Kieffer's (108) referenceto both C. manihot Felt and (Clinodiplosis) brasiliensis Riibs.Houard does not otherwise mention the gall of C. manihot.This point as to whether there are two or only one speciesinvolved cannot be settled definitely until the type specimenshave been compared. It would help, however, to comparerecent specimens of J. brasiliensis with Felt's type of C. manihot,which is still available.COCCINIA INDICA (Naud.) Wight & Arn.The fruit of this plant, which belongs to the Gucurbitaceae,is eaten as a vegetable when green and as a fruit when ripein India.One gall midge has been described from it, namely, Neolasiopteracephalandrae Mani. The female and galls were firstdescribed and figured (127); later, the same authority (128)described the male. The following notes on its life history aretaken from Mani (127). Numerous eggs are laid close togetheron the young vines. The larvae hatch on the fourth day and,penetrating the stem, bore tunnels up and down in the medulla.Yellowish-white o_r pale-green galls, 25-40 mm. x 15-30 mm.develop on the stems. The larvae develop in the galls, reachingmaturity in four weeks. Then they pupate just below the

II M«

e «•tU))_If^1r~iSs:5 MVo/. /. To face page 35

Gall Midges of Economic importance 35while one also causes serious damage to the leaves and thewhole plant, including the root when swedes are attacked.The three midges are Contarinia nasturtii (KiefFer), Dasyneurabrassicae CVVinnertz) and Gephyraulus raphanistri (Kieffer). Theyare dealt with below. A fourth species, whose pale orangeyellowlarvae live in the blossom oi Nasturtium species, is worthyof mention, since it may be found on cultivated Crucifers inthe future. It is only on record so far from Dalmatia and wasdescribed by Rubsaamen (175) as D. nasturtii.Contarinia nasturtii (Kieffer, 1888), the Swede Midge,*syn. torquensyLai&ce, 1906,/lerntawa Rubsaamen, iî^, geisenheyneriRubsaamen, 1917.DIAGNOSTIC CHARACTER. Yellowish "jumping" larvae inseveral types of damage on cabbages (e.g. broccoU, cauliflowers,cow cabbage, kale and savoys), swedes, turnips, rape andradish; flowers swollen, remaining closed; heart leaves crinkledand crumpled, "crumpled-leaf", "curly-leaf" disease; youngshoots and leaf-stalks swollen, distorted and twisted; leadingshoot dead; "many-necked" condition of swedes and turnips,i.e. growth of secondary shoots; subsequent rotting of crown,neck and bulb of swedes and turnips. The "crumpled-leaf"and "many-necked" conditions in swedes are known in Englandcomprehensively as "cabbage top"."]"DAMAGE. The effect of an attack on the flowers is seriouswhen the plant is being grown for seed, as little or no seed isformed (163).The effect of such an attack as "crumpled-leaf" and "manynecked"condition on the well-being and appearance of theplant is obvious. The actual effect on the final weight of thecrop when a root crop is involved is not so well understood.The root may be expected to be lighter than normal. In fact,Taylor (204) found that infested plants produced on the average* Other popular names include Krauselgallmucke, Krusesyge-Galmyggen, laCecidomyie du chou-fleur.t "Many-necked" condition in swedes and turnips is also caused by 6ther insectsand by rodents, i.e. by anything which destroys the terminal bud; "crown-rot" isa bacterial disease; and damage by Thrips tabaci Lind. on cauliflower is not visibletill the outer heart leaves are opened, when a smooth scar instead of the heart

36 Gall Midges of Economic Importanceroots of about half the weight of those from healthy plants.He and Dry (52) were of thq opinion that it was very unlikelythat the midge attack would ever kill young swedes outrightand so necessitate re-sowing.The subsequent rot that usually sets in after midge attackon swedes is probably more injurious than the midge attackitself. This bacterial rotting has been mentioned by severalworkers as a secondary symptom, especially in damp weather,e.g. Walton (221). Davies (44) found on negative evidence,however, that there was no direct relationship between midgeinfestations and the incidence of bacterial rot {Ps. destructans)and dry rot {Phoma Lingam). But Leefmans (117) said that theincidence of attack was correlated with fungal infection inHolland during 1937.T-bivma? (2:12) has described ajid i]]ustrated the generalsymptoms of attack on swedes, turnips, rape, kale, savoys andcow cabbage, which are the more important cruciferous cropsin Devon. In the same paper he has pointed out a curiousdifference in the distribution of attack on cow cabbage andswedes. The heaviest attacks on cow cabbage occurred in thenorth-west of the county, whereas only slight attacks wereobserved in the south and east, although there was a considerableacreage. The opposite was true of the attacks on swedes.Thomas makes two tentative suggestions to account for theobserved facts: (i) that there are biological races of the midgespecific to the different host plants, and (2) that variations inclimate and farming practice in the two areas are responsible.Thomas is continuing his investigations.Mesnil (139) when dealing with damage to cauHflowerstated that, in addition to the complete destruction of theinflorescence, i.e. blindness, two other forms of injury mayoccur. If the cauliflower is attacked about a fortnight afterplanting out, a partial and irregular inflorescence is produced.If it is attacked after the inflorescence is formed,* the larvaedevelop in the flower without causing malformation, bacterialrot sometimes intervening.FLUCTUATIONS IN NUMBERS./ Fortunately the intensity ofinfestation varies grea'tly from time to time. About 1908-14

Gall Midges of Economic Importance 37there was an outbreak on swedes, and to a less extent onturnips, in Yorkshire (52, 204). In 1909 there was a bad attackon swedes at Dundalk, Co. Louth (40). In 1918-20 seriousoutbreaks on swedes occurred in Denmark (160) and later oncauliflowers (72). Gleisburg (78) stated that there were seriouslosses in Upper Silesia, so much so that the cultivation ofcauliflowers had been given up in many instances. He alsoreferred to bad outbreaks in Holland in 1897-1901. WhileFrickhinger (75) stated that Noll, Roesler & Brenner madetheir observations (147) during an outbreak on cauliflower inSaxony. Later Rostrup (161) stated that it was never veryimportant in Denmark until 1926-27, when all the swedes andcabbages in East Jutland and the Islands were infested andbacterial disease followed. Rostrup & Thomsen (162, 163) citeone instance where only i per cent, of the cauliflowers wereMTiattackcd ivi 1927 on oiae. of the. Islands. In ^§^6 also, Walton(221) reported serious damage to swedes and early sown rapein North Wales. During the years 1926 to 1931 there wereserious outbreaks on cauliflowers in north-eastern France, andalso to cabbages and swedes in the neighbourhood of Paris in1926 (148). In Germany cabbages were being attacked toa serious extent in 1931.Dry (52) studied the fluctuations during the latter part ofthe Yorkshire outbreak mentioned above. He found that themidge population increased enormously in the hot summer of1911 and that the first generation of 1912 was very large.Later in the year the summer was particularly wet and cold,and the larvae became scarce. In 1913 the numbers increasedslowly, and again, but more so, in 1914 each successive generationwas larger than the previous one. Dry worked out a swedemidge index figure for the three generations per year from1912 to 1914. This was based on the percentage of plantsshowing the "crumpled-leaf" condition. He found that the increasein population* in a summer depends roughly on the averagemean temperature or on the number of hours of sunshine.The swede midge index figure in different districts wasdetermined by the average date of sowing—the earlier sowingsbeing more heavily attacked—and the distance from the sea.

38 Gall Midges of Economic Importancethe midge being less numerous in the seaside districts than inthe more inland ones. It used to be the opinion among certainentomologists in England 'that coastal areas were more subjectto outbreaks than inland ones, thus contradicting Dry'sobservations. M. Thomsen {in HtL, 20.10.26), on the otherhand, stated that he had noticed no difference in attack eithernear the sea or further inland. Dry reported a definite movementof the adult midges from infested fields to previouslyuninfested ones, while Ritzema Bos (156) and Gleisburg (78)stated that the midges preferred sheltered positions to openones, Thomas (212) found that the intensity of attack wasconsiderably greater in the headlands than in the centres offields of the five crops he had examined, i.e. cow cabbage,savoys, swedes, turnips and rape. Olombel (148) stated that1930 with, its particularly wet summer was a peak year ofmidge increase.It can be seen from Dry's statement of a huge increase in thehot summer of 1911 and Olombel's observation that the underlyingcauses of outbreaks are not well understood. Noll, Roesler& Brenner (147) suggested that the probable cause of outbreaksis a combination of high temperatures and high humidities ata time when there are many larvae in the soil. It does seem,however, from the records in economic literature that theremay be a somewhat regular rhythm of peaks and troughs.A long-term study might well repay its initial monotony afterthe first five years or so.DESCRIPTION AND SYNONYMY. Kieffer (95) first describedthis species, giving descriptions of both sexes, the larva andthe damage, from swollen flowers of Marsh Watercress{Nasturtium palmtre DC). Later Kieffer (99) recorded it fromthe blossom of rape {Brassica napus). It is a typical mediumsizedyellowish-brown Contarinia, and the full-grown larvae arelemon-yellow, while immature ones are yellowish-white.Taylor's (204) illustrations of all stages are reproduced inPis. 2-5.When this investigator discovered that "cabbage top" inswedes, i.e. the "many-necked" and "crumpled leaf" conditions,was caused by a Contarinia species, Kieffer confirmed that

Gall Midges of Economic Importance 39the species involved was C. nasturtii. This alternate type ofdamage is the one that has usually been associated withC. nasturtii in England, and the originally-described flowerdamage has often been forgotten in connection with C. nasturtii.Meijere (134) showed that blindness and distortion of thehearts in savoy cabbages in Holland was caused by a midgeto which he gave the name C. torquens. Both sexes of the midge,the larva and the damage were described by this investigator(134). The larvae live in the leaf axils and are responsible forthe "Draaihartigheid" and the "Krauselkrankheit" in Hollandand Denmark and "Drehherzigkrankheit" condition in Germany,which is exactly the damage that Taylor showed wasdue to the presence of C. nasturtii larvae. In fact, Theobald (209)stated that Schoyen's report in 1910 of C torquens in Norwayreferred undoubtedly to the same insect as that which Carpenter(40) and Taylor (204) reported on as C. nasturtii, butthat he was unable to say which name was the correct one.Laler, following Meijere, other workers in Germany andFrance became accustomed to using this name of C. torquensfor the gall midge causing similar damage on other Brassica.Hahne (79) stated that the damage was usually considered inEurope to be due to C. torquens but that similar damage inEngland has been ascribed to C. nasturtii.P. Bovien, however {in litt. 22.2.29), being of the opinionthat the damage to cabbages and swedes in Germany andassigned to C. torquens was very like that caused by C. nasturtiiin Denmark, suggested that C nasturtii was responsible for alot of the damage* ascribed to C. torquens in north Germany.In 1928 the view was expressed to J. C. F. Fryer by the writer{in litt. 21.3.28) that C. torquens was a biological race of C.nasturtii. Later it was considered that these two species, as wellas C. geisenheyneri Riibs., which is dealt with below, weresynonymous, and this view was communicated to several continentalworkers, e.g. P. A. Blijdorp in Holland, H. Sachtlebenin Germany and P. Bovien in Denmark. Finally, Balachowsky& Mesnil (8) give C. torquens Meijere as a synonym of C.nasturtii Kieffer and state that this synonymy is admitted byseveral entomologists.

40 Gall Midges of Economic ImportanceRiibsaamen found yellowish-white Contarinia larvae livingin the swollen leaf axils of jyoung cabbage plants at Coblenz.Having examined the type female of C. torquens Meijere,Riibsaamen came to. the conclusion that the species he hadreared was a distinct species and named it C. perniciosa (173).He described both sexes, contrasting the female with that ofC. torquens. Some years later he revised his opinion, and inRubsaamen & Hedicke (178) C. perniciosa Riibsaamen is listedas a synonym of C. torquens Meijere.In 1917 Riibsaamen (176) described both sexes and thelarva of C. geisenheyneri, stating that the yellow larvae lived inthe blossom of various Brassica species in Germany. In 1927-28M. Thomsen submitted to the writer for identification midgesthat had been reared in Denmark from larvae living in malformedflowers of swedes grown for seed. These were identifiedas C. geisenheyneri Riibs., but the opinion was then expressed{in lift. 11.1.28) that C. geisenheyneri was synonymous withC. nasturtii Kieffer (162, 163). This view is supported 'byexperimental evidence obtained about the same date byP. Bovien {in Hit. 19 and 23.4.38) in Denmark. He placedseveral hundreds of C. nasturtii reared from "crumpled-leaf" onswedes in a cage with flowering swedes and found that theirGi produced exactly ^the same flower malformation that isgenerally ascribed to geisenheyneri.It seems, therefore, that C. torquens Meijere, C. perniciosaRiibsaamen and C. geisenheyneri Riibsaamen are best consideredas synonyms of C. nasturtii Kieffer.It would, however, be interesting to attempt to transferC. nasturtii reared from Nasturtium palustre to cultivated Brassica,thus making quite sure that Kieffer was correct in his determination(99) that C. nasturtii on Brassica was the same speciesthat he had found earlier (95) on various species oi Nasturtium.If this transference proved' impossible, the name C. nasturtiiwould have to be reserved for the midge living on Nasturtiumspp., while C. torquens Meijere (syn. perniciosa and geisenheyneri)would have to be considered as the name of the separatespecies living on Brassica spp.Other species have beeli described from Brassica flowers by

Gall Midges of Economic Importance 41Tavares, and it is possible that they also will prove to besynonymous with C. nasturtii. They will be mentioned againin the section CLOSELY ALLIED SPECIES (p. 49):DISTRIBUTION. This species is a common European speciesbeing recorded as follows: as C. nasturtii in Denmark (159),France (95, 151), Germany (120), Sicily (190), Italy (215),Portugal (196), Belgium (138) and Great Britain. In the lastnamedcountry, Carpenter (39, 40) recorded it from Dundalk,Co. Louth, in Ireland; Cameron (38) reported it from Scotland,Walton (221) from North Wales, and it is on record frommost countries in England.As C. torquens it is on record from Holland (134), Norway(181), Germany (78) and France (130).As C. perniciosa it has been recorded from Germany (173)and as C. geisenheyneri it has been reported from Germany (176)and Norway (162).LIFE HISTORY, The biology of this midge on plants ofeconomic importance has been studied in England by Taylor(204), Dry (52) and Thomas (212), in France by Olombel (148)and Mesnil (139), in Denmark by Rostrup (160, 161), inHolland by Spithost (189) and Leefmans (116, 117), and inGermany by Roesler (157) and Noll, Roesler & Brenner (147).The eggs are 0-27 mm. x o-o8 mm. with a pedicel measuringo-o6 mm., according to an observation of P. Marchal quotedby Olombel. They are laid on the youngest parts of the plant,both the flower-buds and the leaves in the centre accordingto the stage of growth of the plants. When the leaves are chosenthe eggs are laid in strings and clusters, mostly near the baseof the leaf-stalks on their upper surface, while some are actuallyplaced on the leaf blades. About 15-20 eggs are depositedtogether, and one female will lay several (5-6) batches.Leefmans in Holland found that each female matured 66-124eggs and laid them in batches of 4-48 on at least 3-5 plants.The same authority stated that the females are very inactiveat temperatures below 68° F.The larvae hatch in about 3-4 days (4-5 days at 68° F. or9 days at lower temperatures according to Noll, Roesler &Brenner, and 3-5 days in summer according to Leefmans).

42 Gait Midges of Economic importance .They live on almost any part of the plant, Olombel statingthat mature larvae have even been, found in quite well-• developed inflorescences of cauliflowers, Taylor stated thatthey feed on the juicy tissues by gnawing. On the other hand,Olombel is more correct when he says that they do not appearto gnaw. He suggests that their digestive juices exercise achemical action on the plant provoking an exudation of aliquid in and on which the larvae live.As the result of their presence after a few days (3-4 on swedesaccording to Taylor,' 4-5 on cauliflowers according to Mesniland 9-11 on cauliflowers according to Olombel) the typicalsymptoms of attack begin to appear. These have been ascribedto a checking of growth at the points of attack and the secretionof toxins by the larvae. The stalks become swollen and, bendingsharply inwards across the top of the plant, press upon andcompact the terminal bud. In other cases the leaves start tobecome crumpled.The number of larvae present naturally influences theintensity of these symptoms, but in any case the unfolding ofthe terminal bud is delayed, thus affecting the growth of theplant as a whole. The larvae are soon protected from spraying.According to Taylor the feeding period In Yorkshire lastsabout three weeks, and in France Mesnil stated 10-12 dayswere sufficient, while Olombel said it was about 15 days.In Holland, Leefmans found that in 1936 the larval stagelasted 15-24 days in summer, while Noll, Roesler & Brennerstated that in Germany 68°~7i-6° F. represented the optimumtemperature and development was then completed in 7-8 days.According to Olombel the larvae can only live in an almostliquid medium and, although they can remain in water severaldays, drought has an immediate lethal effect on them. Hestated in support that the wettest ground good for the productionof cauliflowers is the most subject to attacks by this midge,and the wet summer of 1930 was a peak year in midge increase,Leefmans found in submersion tests that normal adults emergedafter larvae had been kept under water for 3 months. Pupationwas delayed.When fully grown the laryae are a!bout -j^th inch in length,

Fig. 2.—Pupa of Swede Midge. Magnified 45 times. (After T. H. Taylor)

Gali Midges of Economic Importance 43pale yellow in colour (they are more white than yellow in theimmature stages) and can "jump" like other Contarinia species.They "jump" from the plant to the soil where pupation takesplace in silken cocoons covered with particles of soil in theupper 2-inch layer of soil. After this movement has takenplace it is still often possible to diagnose the midge attack bythe presence of damaged plants, although the unpractised eyemay confuse it with that of other insects.In the summer generations the pupal period varies fromabout 10-13 days in France, 6-7 days in Holland, to about2-3 weeks in Yorkshire. This period, of course, depends onthe local and seasonal conditions. The last generation of theyear passes the winter in the soil.Mesnil suggested that the larvae overwinter and pupationdoes not take place till the spring, particularly as the firstemergence is relatively late in the year. Leefmans stated thatthe larvae hibernated in their cocoons and pupated in thespring, and added that some midges emerged in July, 1937,from pots of soil in which the larvae had pupated in July, 1936,although some had emerged in August, 1936. Noll, Roesler &Brenner state definitely that the larvae of the third generationand also some of those of the first and second generation overwinterand pupate in the spring (see C. pisi, p. 71). When theinsect is ready to emerge, the pupa works its way out of thecocoon and moves up to the surface of the soil until the frontportion protrudes into the air. Emergence of the midge thentakes place.This species has several overlapping generations a year.In France three generations have been recorded. In EnglandTaylor and Dry found that three was the usual number, butthat occasionally there were four, as in the hot summer of 1911.Rostrup found that the same was true of the midge in Denmark,a partial fourth generation occurring in some places, andMeijere found the same in Holland. Roesler stated there werefive generations in Germany, the fifth being very scanty. Onthe average, however, there were three generations a year.It is generally- accepted that second and third generationsusually do most harm.

44 Gall Midges of Economic importanceIn Yorkshire, Taylor recorded that the four generations in1911 appeared in June, July, August and September, eachtaking just less than six weeks. Taylor reckoned the first fortnightin June was the average date for the beginning of theswede midge season. This is the tirhe when May-sown swedeswould be ready for attack. However, he mentioned the possibilityof an earlier generation on some other host plant thanswedes. Theobald (207) reported receiving larvae from CountyLouth which descended from the plants to the soil onMay 10-14, which is very early, and again on July 15.In north-east France, Olombel also found the midges firstemerging in the first ten days of June, injured plants beingnumerous by June 16-21, the first one being recorded onJune 10. Adult midges of the first generation appeared betweenJuly 6 and 8 and those of the second on August 9-11. Thelarvae of the third generation left the plants for the soil inlate August and early September. Olombel was dealing withthe swede midge on cauliflower. Mesnil never found anadult in the open before June i, but in frames where thetemperature was slightly higher they started to appear aboutMay 15.In Holland there were three main flights of midges in 1936about June 20-27, July 17-21 and August 24, while in 1937there were only two peaks, about July i and July 30-31. Thenumbers of midges were obtained by the use of emergencecages in the field. These had wooden sides and wire gauzetops. These 'latter could be darkened by covers. The midgeswere collected in a glass tube inserted in one side.Rostrup (161) stated that in Denmark the attack started in1926 at the end of May or beginning of June, whereas in 1927it only began about mid-July. In 1926 the first-generationlarvae were present from the end of May to the end of June,those of the second in July, and those of the third in Augustand September. Those of the partial fourth generation whichoccurred in some places were present in late September andpossibly in October. Although no general statement could bemade in regard to the effect of the date ,df sowing, Rostrupstates that in the majority of instances the crops sown early

Gail Midges of Economic importance 45(before April 25) seem to have been most resistant (sic) to theattack.Roesler stated that in Germany the generations occur atsuch remarkably regular dates that a spray calendar can begiven (see p. 48).FOOD PLANTS. These include many cultivated Brassica spp.,including swede [B. rutabaga), turnip [B. campestris L.), rape {B.napus L.) and cabbage {B. oleracea L.), and broccoli, cauliflower,cow cabbage, kale and savoy. The midge is also recorded fromB. amplexicaulis Janka (190) and charlock or wild mustard{Sinapis arvensis h.) (52, 102).It is also reported from various species of Raphanus, e.g. wildradish [R. raphanistrum L.) (216) and R. caudatus L. (97).Frickhinger (75) includes Cochlearia armoracia L. (horseradish)in his list of host plants.Riibsaamen & Hedicke (178) list Nasturtium palustre, jV". silvestre,JV. officinale, jV". amphibium, Raphanus sativus and R.raphanistrum as host plants of Contarinia nasturtii; the leaves ofBrassica spp. as those of C. torqmns {C. perniciosa); and theblossom of Brassica spp. as those of C. geisenheyneri.It was originally described from marsh watercress [Nasturtiumpalustre DC.) (95) and has also been recorded from creepingwatercress {N. silvestre R. Brown) (102, 215), and, with a queryas to the correctness of the identification of the species of theContarinia, from JV. pyrenaicum R. Brown (85, 151).It is quite possible that the species found so commonly onBrassica and Raphanus spp. is not the same as that originallyfound on Nasturtium. Dry (52) was of the opinion that wildNasturtium species were not of much importance in determiningthe abundance of the midge on swedes in Yorkshire.RESISTANT VARIETIES. Mesnil (139) found that all thevarieties of cauliflowers grown in north-east France wereequally susceptible to attack. Spithost (189) also reported thatno varieties of cabbages were immune, but that the period ofsusceptibility diflPered with the variety. Regarding variety ofswede, Walton (221) found severe attacks on Balmoral, Bangholm,Best of All, Buffalo, Dreadnought, Echpse, Lord Derby,Magnum Bonum, Model, Monarch and Superlative. He came

46 Gall Midges of Economic Importanceto the conclusion that variety had little effect on infestation.Davies (44), however, found that Wilhelmsberger sufferedleast from attack.It has been recorded on occasion that turnips are notattacked as much as swedes, e.g. Walton (221) stated thatturnips even when sown next to damaged swedes wereunattacked. However, it appears probable that this is not aquestion of plant resistance or preference but rather due toa difference in time of development of the susceptible stagesof the plants in relation to the appearance of the adult midges.Turnips are normally sown much later than swedes.NATURAL ENEMIES. Dry (52) records an Empid fly suckingthe juices of the midges, and states that small spiders alsoaccounted for some.The same investigator reared a number of Proctotrupidswhich parasitize the larvae. With this exception no parasiteshave as yet been recorded.CONTROL. Several methods of cultural control may beemployed to prevent the midge when present in small numbersfrom reaching pest proportions. Rotation of crops, if accompaniedby clean cultivation where possible, is quite effective,as the midges will move away to other areas on the one hand,and on the other will not travel very far from an old Brassicafield into a new one.In order to thwart the migrating midges, trap crops or stripsof swedes, e.g. in corn headlands, may be planted and, wheninfested, removed and destroyed. Taylor (204) when trying thismethod found it successful, but pointed out that migratingmidges oviposit on the first available plant which theyencounter. For this reason he stresses the danger of allowingswede-remains to develop. Care should be taken to remove allthese. In addition, wild host plants may serve as usefulreservoirs if allowed to remain.As the symptoms^ of attack are easy to recognize, it mightalso pay to lift and burn at the beginning of the season anyyoung plants showing the typical damage, especially with acauliflower crop. It is not sufficient to hoe out the plantswithout removing them as the larvae might often be able to

Gall Midges of Economic importance 47complete their development even if only undersized midgeswere the result.Further, as the larvae are very susceptible to drought, discriminatehoeings to dry up the surface of the soil should bemost useful if these operations were timed correctly. Thecorrect periods would be the occasions when the larvae havejust left the plants for the soil. The exact dates would have tobe worked out in each particular area and sometimes accordingto the season. Noll, Roesler & Brenner (147) argued that increasedcultivation had little effect.Finally, it is generally accepted that the earHest sowings aremost heavily attacked, and thus a delay in sowing might beadvantageous.The use of insecticides in practice is restricted to the eggand very young larval stages and the fully-grown larval stagein Hat soi\. Tbe adxilt tradgcs aie too sbort Vwed aad \i\^ larvae,soon gain good protection from sprays by the malformation ofthe plants.Little success has accompanied attempts to kill the larvae in. the soil. The only reported success is that of Olombel (148),who claimed that cauUflowers grown on soil treated with1,000-1,300 lb. sylvinite or 1,300 lb. paradichlorobenzene peracre were less severely attacked. According to Balachowsky &Mesnil (8), however, Vayssiere attempted without success tokill the larvae in the soil by the use of paradichlorobenzene.In addition, various workers have tried without success toreduce the numbers of the larvae in the soil by means ofartificial manures (e.g. 139), but Gleisburg (78) stated thatareas treated with lime suffered least and Noll, Roesler &Brenner (147) reported that some mortality was caused by theapplication of calcium cyanamide and kainit. Davies (44) alsowas successful in reducing loss from midge attack by theappUcation of nitrate of soda at the rate of i cwt. per acre,provided it was appUed in time for the young plant to makeuse of it, i.e. putting it in the drills either at the time of sowingor just before.Repellents have been tried out against the adult midges toprevent them egg-laying. Mesnil (137) reports tbat of seventeen

48 Gail Midges of Economic Importancesubstances tested creosote or naphthalene and pyridine werepromising. Each substance was mixed with ground nut oil(1:9), the mixture afterwards' being emulsified in water atthe rate of 2 per cent, by means of ammonium oleate. The oilwas added to retain the» odour 1 of the repellent and also as asolvent and spreader. Para-phenylenediamine, ethyl hydrosulphide,ammonium valerianate and chloronaphthalenewere also used in ground nut oil and emulsified in water withammonium oleate (2), but were all more effective as contactinsecticides than as repellents, and at lethal concentrations thefoliage was scorched. Later, Mesnil (139) reported that thesuccess at first attributed to such repellents had been found tobe due to the insecticidal effect of the ground nut oil. Noll,Roesler & Brenner (147) also failed to discover an effectiverepellent.The use of sprays to kill the eggs and very young larvae has,however, been successful. Gleisburg (78) mentions that inHolland spraying once a week with 6 parts by weight oftobacco leaves in 100 parts of water, from the first seedlingstage onwards, had proved successful. Spithost (189) said thatthe best results had been obtained by spraying weekly with asolution of 2 per cent, soap and i j>er cent, methylated spirit.He also recorded that on a small scale weekly dusting withnaphthalene in fine flakes was effective.However, nicotine sprays appear to be best, and detailedwork on these has been carried out in Germany, France and•Holland. Roesler (157) found o-i-o-i5 per cent, crude nicotineand 0-5 per cent, soft soap preferable to derris or pyrethrum.The spray was directed from above into the hearts of the plants.This investigator found that the appearance of the generationsin Germany was so regular that he could make a spray calendar.Spraying was recommended against the first generation onMay 25 and June 22; against the second generation on June 12,18 and 24; against the third on July 9, 15 and 24 and sometimesa fourth application; against the fourth on August 12,19 and 25. Roesler said that no spraying was necessary againstthe fifth generation and only occasionally against the first andfourth. It is, however, considered a short-sighted policy not to

Gall Midges of Economic importance 49spray the first generation and so allow a second generation todevelop without any such setback. Roesler also pointed outthat seedlings in frames are subject to attack, and he suggestedspraying such plants at 8-day intervals throughout the seasonfrom April onwards.Mesnil (138) went thoroughly into the problem in northernFrance m 1935-36. Closely following ordinary commerciaZpractice, he had three sowings of cauliflowers followed by threetransplantings on May 11 and 25 and June 6. Harvesting ofthese plants began on July 21. Three sprays were tested1-2 parts nicotine sulphate to 5 parts of sodium sulphoricinate;I part pure nicotine to 5 parts of sulphoricinate; and i partnicotine, 7-5 parts oleic acid and 5 parts of commercial volatilealkali. Each of these was in i ,000 parts of water. The sprayswere applied by portable apparatus twice a week from May 13to July 16 at the rate of 11 pints to 30 plants of average size.The results were wholly successful, the unsprayed controlsshowing 77-83 per cent, injury, while those sprayed sufferednot more than 0-39 per cent, and with these the damage wassustained before transplanting. The cost was 2^ per cent, ofthe market value of the plants. In practice, Mesnil suggestsnot more than 10 applications at slightly longer intervals.Leefmans (117), after a considerable number of trials inHolland, recommended o-i per cent, nicotine and i| per cent,soap for headed cabbage and 2 per cent pyridine and 1} percent, soap for cauliflower. He said that headed cabbage shouldbe sprayed twice a week, while slight infestations on cauliflowermight be controlled by weekly applications. The same investigatorstated that derris could be recommended as a dustinsecticide, but spraying with a strong jet is more eff"ectiveowing to the larvae and eggs being washed away.CLOSELY ALLIED SPECIES. Contarinia tudensis Tavares, femaleonly described, bred from galled flowers of Erucastrum incanumKoch, at Tuy, Galicia, Spain (197).C. pontevedrensis Tavares, male and female described, rearedfrom larvae living in the flowers of Raphams silvestris Lam. andBrassica napus L. near Pontevedra, GaUcia (198).C. gallaica Tavares, male and female described, reared from

50 Gall Midges of Economic Importancelarvae living with C. pontevedrerisis in the flowers of R. silvestrisLam. and B. napus L. at Tuy and near Pontevedra, Galicia (198).The galls of the above three species are stated to be indistinguishablefrom those of Gephyraulus raphanistri Kieffer.In addition, the writer has in his collection specimens ofmales and females of a species which is exceedingly similar toC. pontevedrerisis. They were received in 1938 and 1939 fromR. Dieuzeide, who reared them from galled flowers of B. fruticulosaCyrill spp. radicata Dsf. in Algeria.MATERIAL. C. nasturtii. Males and females, England, Yorkshire(reared from leaves of swedes, part of T. H. Taylor'smaterial, 4th generation 1911), Kent (reared from leaves ofturnips) and Devon (reared from leaves of swedes by D. C.Thomas).C. torquens. Male and female, France (received from P.Marchal, reared 1926).C. geisenheyneri. Male and females, Denmark (received fromM. Thomsen, reared from flowers of swedes, Jutland, July,1926).MOST IMPORTANTREFERENCESKIEFFER, J. J. 1888. Ent. Machr., 14, 263-4 {Diplosis nasturtii)MEIJERE, J. C. H. DE. 1906. Tijdschr. Ent., 49, 18-24, 28 and pi. 3 {Contariniatorquens) "RuBSAAMEN, E. H. 1914. MarcelUa, 13, 111-13 [C. perniciosa)RiJBSAAMEN, E. H. 1917. S.B. Ges. naturf. Fr. BerL, 92 (C. geisenheyneri)TAYLOR, T. H. 1912. Univ. Leeds and York. Counc. agric. Educ, no. 82,21 pp. (C nasturtii)DRY, F. W. 1915. Ann. appl. Biol., 2, 81-108 (C nasturtii)GLEISBURG, W. 1920. Dtsch. landw. Pr., 47, 705-6 (C. torquens)ROSTRUP, S. 1921. Mord. JordbrForskn., 3-4, 301-12 (C. nasturtii)WALTON, C. L. 1927. J. Minist. Agric, 34, 547-51 (C. nasturtii)ROSTRUP, S. 1928. Tidsskr. Planteavl., 34, 692-716 (C nasturtii)SPITHOST, C. 1929. Versl. Pl.îekt. Dienst Wageningen, no. 54, 24 pp. (C.torquens)DAVIES, W. M. 1931. Welsh J. Agric., 7, 319-32 (C. nasturtii)OLOMBEL, M. 1931. C.R. Acad. Agric. Fr., 17, 178-82 (C. torquens)ROESLER, R. 1937. Kranke Pflanze, 14, 124-9 C-^- torquens)MESNIL, L. 1937. Rev. Path, veg., 24, 70-7 (C nasturtii)LEEFMANS, S. 1937. Meded. Tuinb.-Voorlichtihgsdienst, no. I, 28 pp. (C.nasturtii)^

Gall Midges of Econo.mic Importance 51LEEFMANS, S. 1938. Meded. Tuinb.-Voorlichtingsdienst, no. 5, 42 pp. (C.nasturtii)MESNIL, L. 1938. Ann. Epiphy. Phytogen., 4, 281-311 (C. torquens = nasturtii)* NOLL, J., ROESLER, R. & BRENNER, J. 1942. Arb. physiol. angew. Ent.Beri, 9, 1-44 (C nasturtii)FRICKHINGER, H. W. 1943. Kranke Pflanze, 20, 65-8 (C. nasturtii) (review ofabove).Dasyneura brassicae (Winnertz, 1853), the Brassica Pod Midge.tDIAGNOSTIC CHARACTER. Swollen and prematurely ripeningand yellowing siliquas of cabbages, swedes, turnips, colza orrape and radish. Known as "bladder pod" in Cambridgeshire.DAMAGE. Normal development of seed is prevented, consequentlysevere infestations result in considerable loss to theseed harvest. This may be either of two types, seed for sowingor seed for the oil, e.g. rape seed or rape cake. Such attackshave occurred in Denmark (1919 and 1920) on cabbages andturnips (71, 72) and in Germany on summer crops of oilseedproducingcrucifers, especially experimental sowings of summerrape (188).DESCRIPTION. Winnertz (225) originally described both sexesof this species, which is a typical medium-sized Dasyneura, witha red abdomen in the female. The larvae are gregarious, whiteand "non-jumping".DISTRIBUTION. Widely distributed in Europe, being on'record from Denmark (121), Norway (182), Holland (155),Germany (187, 188, 225), Poland (227), Bessarabia (i),Czecho-Slovakia (25), France (115) and Great Britain. Inaddition, Linnaniemi (122) has recorded it from Finland.In Great Britain, Theobald (205) reported it from RomneyMarsh and Thanet, and Bagnall & Harrison (3) recorded itin Durham. It also occurs commonly in Cambridgeshire andLincolnshire, while the writer has found it in Hertfordshire.Probably it occurs throughout England.. LIFE HISTORY. The chief biological studies of this species are* Not seen in original.t More often called the Pod Midge or Bladder Pod Midge. Other popular namesinclude the Turnip and Cabbage Seed Midge, Kohlgallmiicke, Kohlschotentniickeand Koolzaadgalmug.

52 Gall Midges of Economic Importancethose of Speyer (187, 188), but a complete detailed study isstill required to clear up several points.Laboulbene (115) stated that the fenaale pierces the siliqua inorder to insert her eggs, but tllis is extremely unlikely owingto the structure of the ovipositor. Speyer (187), on the otherhand, records watching them oviposit repeatedly through thepuncture holes made by weevils {Ceuthonhynchus assimilis. Payk.and allied species). In fact, later (188), he states that this midgeis only dangerous to summer crops of crucifers where C.assimilis is present. However, it seems improbable that the midgeis entirely dependent on weevil punctures for successfuloviposition. This question is at present being re-investigated insouth-east England.The larvae live gregariously inside the siliquas (PI. 6,Fig. I') but it is not quite certain in what manner exactly theyfeed. Laboulbene {loc. cit.), states that, by sucking, they forma humid exudation, which lines the siliqua. It is highly probablethat they scratch the inside surface and also sometimes thesurface of the developing seeds, and then feed on the exudingsap. The presence of shrivelled seeds, which are sometimesfound in infested siliquas, supports this view. At any rate, as aresult, the siliqua often becomes malformed and unshapely.This in itself is an indication of differential growth which is anatural sequence of rasping of the internal tissues.When the siliquas are not badly misshapen, it is likely thatthe larvae have been feeding more on the developing seeds thanon the siliquas themselves. In any event, normal seed productionis prevented, although often odd seeds develop. Infestedsiliquas are discoloured and often turn yellow, giving theappearance of premature ripening. Varying numbers of larvaeper fruit have been reported, e.g. Laboulbene, 15-20 or more;Theobald, 35; Ritzema Bos, 50; and Winnertz, 50-60.The larvae, which are white, are full grown in about fourweeks, and then, when the siliquas open, descend to the soil.They do not "jump" as is the habit oi Contarinia larvae. Pupationtakes place in a cocoon about |—3| in. deep in the soilaccording to Speyer. Theobald (20^) stated that the adultmidges appear a fortnight later, while the author has reared

Gall Midges of Economic Importance 53midges between July 3 and 12 from larvae collected on June 22.The last generation of the year spends the winter as full-grownlarvae in the soil, pupating the following spring.There are several generations a year. In England Theobald(208) found adult midges in May and June, and larvae up tothe first week in October. In Germany Speyer (188) found therewere about six generations a year from May to October, andstated that some of the larvae of the third and later generationsdid not pupate until the following spring.This authority (187) recorded that dampness in conjunctionwith warmth and the presence of Ceuthorrhynchus assimilis arefactors favourable to the increase of this species. It has also beenstated (30) that the presence of even a few of the weevils orallied species is enough to help the first generation to establishitself and increase. Woroniecka (227) mentioned that rainyweather during June delayed the ripening of the pods, soincreasing the damage done by this midge.FOOD PLANTS. Several Brassica species, including rape or colza{B. napus L.) from which it was originally described, cabbage{B. oleracea L.), turnip {B. campestris L.), swede {B. rutabaga),Jersey cabbage {B. cheiranthus Vill.) (93, 98), charlock {^binapisarvensis L.) and black mustard (S. nigra L.) (187). Rape andfield cabbage seem to be most often attacked.It has also been recorded on radish {Rapkanus sativus L.).Rakhmaninov & Virzhikovskaja (153) recorded a midge,which they stated was possibly this species, in the siliquas ofCamelina sativa Cr., in the Tambov Government, Russia. Theauthor has also received an unidentified but very similarDasyneura species from the flowers of C sativa from the Voronezhdistrict, just south of Tambov.Speyer (188) states thiat in threatened areas white mustard(Sinapis alba L.) only should be grown, and that wild charlock{S. arvensis) should be destroyed, thus implying that while thelatter plant is a host, plant, the former is not.An intensive study of the host plant range of the three midgespecies, D. brassicae Winn., Gephyraulus raphanistri Kieffer andC. nasturtii Kieffer, especially as regards wild crucifers, mightbe illuminating and even helpful in their control.

54 Gall Midges of Economic importanceRESISTANT VARIETIES. None has been reported, but thisaspect has not been intensively studied.NATURAL ENEMIES. De Gaulle (,76) listed this midge as a hostof Inostemma boscii Jur. in France,' but this is not an altogethersatisfactory record. |Speyer (188) states that a Proctotrupid Platygaster sp. (? nigraNees) parasitizes the eggs of this midge and that two Chalcids[Pseudotorymus brassicae Ruschka and Tetrastichus brevicornisThorns.) parasitize the larvae. Up to 68 per cent, of the midgelarvae were parasitized by the Chalcids in August, 1921.Certain of the third and later generations of the midge escapebeing attacked by these Chalcids. T. brevicornis disappears inSeptember a month earlier than Pseudotorymus brassicae, althoughthey both appear at the same time in the spring. T. brevicornisprobably has one generation a year less than P. brassicae,two or three generations at the most.Speyer (187) also stated that the larvae are attacked by afungus.INQUILINE. The author has reared an unidentified Clinodiplosisspecies from cabbage siliquas attacked by D. brassicae.CONTROL. NO satisfactory method of controlling this midgehas-yet been recorded, but Rademacher (152) stated that thethen recent outbreaks of the midge in Germany had beenchecked by the enormous decrease in the cultivation ofcruciferous oil plants. He also suggested that they couldbe avoided in future by crop rotation. Summer crops mustnot be grown following winter rape, especially as thepresence of a few weevils has been stated to be of greatimportance in helping the midge to become established inthe spring.Handpicking of infested siliquas might be of some benefit inseed crops of very limited dimensions, but it is not easy to spotevery infested fruit.It has also been suggested that deep ploughing will destroythe overwintering larvae. In fact, nicely-timed intensivecultivation seems to be the most hopeful line of control andworth while developing.If also the midge is dependent on the presence of the weevils,

Gall Midges of Economic Importance 55any successful measures taken against the latter should reducethe numbers of the midge.If it is established that cruciferous weeds can maintain themidge, the eradication of such weeds should be practised.CLOSELY ALLIED SPECIES. There has been some confusion asto the validity of the two species Dasyneura brassicae Winn, andGephyraulus raphanistri KiefTer. Some authorities, e.g. Riibsaamen(170), have thought that they might be identical.Later, however, Riibsaamen (175) recognized that there weretwo distinct species and made raphanistri Kieffer the type of anew genus Gephyraulus, while retaining brassicae Winn, in thegenus Dasyneura. Balachowsky & Mesnil (8) also have confusedthe two species, referring to D. brassicae as "la Cecidomyie desfleurs de Cruciferes" and in dealing with it have made severalother statements that apply not to brassicae but to raphanistri.Actually, although the two species are very similar, when onecompares the females it is obvious, though hard to describe inwords, that there are two distinct species—viz. D. brassicae thelarvae of which live in the siliquas, and G. raphanistri the larvaeof which live in the swollen flowers. The differences, however,are not considered to be of generic magnitude.MATERIAL. Males, England (Hertfordshire) and Germany.Females, Germany. Larvae, England (Hertfordshire).MOST IMPORTANTREFERENCESWiNNERTZ, J. 1853. Linn. Ent, Bed., 8, 231-3 {Cecidomyia)SPEYER, W. 1921. Mitt. biol. Anst. (Reichsanst.) Berl, 208-17 {Dasyneura)SPEYER, W. 1923. Arb. biol. Abt. {Anst.-Reichsanst.) Bed., 95-108.Gephyraulus raphanistri (Kieffer, 1886), the Brassica Flower Midge.*DIAGNOSTIC CHARACTER. Flowers of cabbages, swedes,turnips, colza and radish swollen and remaining closed, containing"non-jumping" white larvae. The "non-jumping"character of the larvae is the best one for the separation of thelarvae of this species from those of C. nasturtii Kieffer without alens or in the field. The colour difference is not always obvious.DAMAGE. The calyx becomes enlarged, the corolla short, the* Also known as the Colza Flower Midge, Kaalinkukkasaaski.

56 Gall Midges of Economic Importancestamens shortened and swollen, the ovary hypertrophied. Theflowers remain closed and swoUenl No fruit and therefore noseed develops. Hence this midge is apt to cause loss to cropsgrown for seed. It appears seldom,in economic literature, buthas most probably been overlooked or recorded as C. nasturtii,perhaps even as D. brassicae. However, Noel (144) listed it asa pest of radish in France.DESCRIPTION. Kieffer (94) described the male, female, larvaand gall of this species. At first sight it appears to be a mediumsizedDasyneura, the females having yellowish-brown abdomens,but a comparison reveals that it is slightly a-typical of thisgenus, especially as regards the ovipositor. Riibsaamen (175)made it the type of his newly-erected genus Gephyraulus. Thelarvae are white.DISTRIBUTION. Central and western Europe, being recordedfrom Portugal (194, 196), France (141), Holland (136),Germany (165), Finland (89, 122) and England. In the lastnamedcountry, Bagnall & Harrison (3) have reported it fromNorthumberland and Durham. /LIFE HISTORY. Very little has been recorded concerning thelife history of this species, but Tavares (202) described theoviposition. There are at least two generations a year, in thespring and summer. The larvae, which are white and gregarious,leave the flowers in order to pupate in the soil. It isnot stated whether the winter is passed as larvae or as pupaein the soil, but probably it is as fully fed larvae.FOOD PLANTS. Several Brassica spp., including cabbage[B. oleracea L,), colza or rape [B. napus L.), swede {B. rutabaga),turnip {B. campestris L.) and B. adpressa Bois. (195).It has also been recorded on radish [Raphanus sativus L.) (120),wild radish (R. raphanistrum L.), R. caudatus L. (97) and R.silvestre Asch. from which it was originally described.Finally it has been reported from Erucastrum pollichii Schimp.& Spann. by Tavares (194).NATURAL ENEMIES. The following parasites of this midgeon radish are described by Kieffer (109), Synopeas raphanistriKieffer, Prosactogaster jioricola Kieffer and Platygaster cruciferarumKieflfer. • ' -

Gall Midges of Economic Importance 57CONTROL. None has been recommended.CLOSELY ALLIED SPECIES, See D. brassicae Winn, under thisside-heading (p. 55).MATERIAL. Males and females, Germany (on Raphanussativus).MOST IMPORTANTREFERENCEKiEFFER, J. J. 1886. Z- Naturw., 59, 324-6 {Cecidomyia).EGG-PLANT or AUBERGrNE{Solarium melongena L.)Del Guercio (45) described a lateral or terminal globulargall, covered with short hairs caused by an unknownCecidomyid on this plant in Eritrea. The tissue of the gall wasspongy and there was a central cavity in which a yellow larvawas found.The author has in his collection females of an Asphondylia sp.reared by W. V. Harris in June, 1936, from galls on S. melongenaL. found at Morogoro, Tanganyika Territory. The gall wasnot described in the relevant correspondence. This gall midgewas added to the list of injurious insects of Tanganyika byHarris (82).Finally, E. McC. Callan has stated verbally (August, 1944)that a Cecidomyid has been found attacking the Egg-plant inAntigua (B.W.I.). This may be the midge that has been identifiedas C. lycopersici on egg-plant in Hawaii (see p. 79), but it hasnot yet been possible to examine either Callan's or Holdaway'smaterial.LEEK [Allium porrum L.)In the autumn of 1941 the larvae of an undetermined speciesof Clinodiplosis were reported by C. D. R. Dawson as infestingthe seed-heads of leeks grown for seed. Specimens of the larvae,males and females, are in the author's collection.The orange larvae were first noticed when the seed-headswere harvested. They were in the pithy part of the centre of

58 Gall Midges of Economic Importancethe heads where the flower-stalks join on to the main stem. It isthe practice to harvest the seed-hâds at the end of Septemberor early October, tie them in bunches* and hang them up inbarns or glasshouses to dry off and finally ripen. This is completedby Christmas. At the time of harvesting the midge larvaewere apparently causing some of the heads to break up in thefields but not in large numbers. By the beginning of November,however, many of the heads were falling to pieces and some ofthe larvae were in addition found inside the seed capsules.They were thus of appreciable nuisance importance.Gall midges of this genus have frequently been reported inthe seed-heads of many plants, e.g. lobelia, hollyhock, cosmos,gladiolus, lettuce (see p. 59), red clover, etc. (24), but in noinstance has any actual damage to the seeds been seen. Theselarvae seem rather to feed on the tissues comprising the seedcasesand heads, particularly when the latter are drying off andthe seeds are ripening. Certain other species of this genus,however, have other habitats, e.g. C. hydrangeae Barnes, whichcauses blister galls on Hydrangea leaves (23).LENTIL [Lens esculenta Moench)The true lentil is Lens esculenta Moench, which is perhapsthe oldest food plant grown. Noel (143) in a list of enemies oflentils mentions three gall midges, namely Asphondylia erviRiibsaamen, Dasyneura viciae (Kieffer) and an unidentifiedCecidomyid listed by Darboux & Houard (43). There is someuncertainty as to whether Noel was referring to L. esculenta orcertain species of Vicia. V. tetrasperma Moench and V. hirsutaKoch have both been placed in the genus Ervum, which usedto include Ervum lens L. == L. esculenta Moench, and is stillsometimes known popularly as signifying Lentils.If Noel was referring to the correct L. esculenta, it is unknownon what authority he stated that the above-mentioned midgesattack it. It is more likely that he was thinking of lentils as thespecies of Vicia that used to be placed in the genus Ervum. Thusthe food plant of Asphondylia ervi Riibsaamen is Vicia {Ervum)

Gall Midges of Economic importance 59and other plants (see below); while the recorded food plantsof Dasynema viciae (Kieffer) include Vicia [Ervum) tetraspermaMoench; and Darboux & Houard gave Vicia (Ervum) hirsuta asthe food plant of the unidentified gall midge.Riibsaamen (i68) originally described A. ervi from Vicia(Ervum) hirsute from Germany. It is a typical Asphondylia. Thelarvae are yellow and live singly in the pods, which are swollenat the base. Pupation takes place in the pod. Bagnall & Harrison(5) stated that they had found this species in Devon duringSeptember on the same plant species. Felt (67) records receivingwhat is possibly this species from soy bean pods in Japan.Felt, in correspondence (3.12.18) to S. I. Kuwana, stated thatthere ,was little doubt in his mind but that the specimens werethe typical A. ervi Riibsaamen. The author received in 1937similar specimens for identification from H. Yuasa of theImperial Agricultural Experiment Station, Nishigahara, Japan,and replied that he was of the opinion that it should bedescribed as a new species. Males and females of this consignmentare in the author's collection. It has not yet been possibleto compare them with A. ervi reared from V. hirsuta. Yuasa &Kumazawa (228) gave the distribution of this Asphondylia sp.in Japan and listed the following Leguminous plants. Glycineussuriensis, Cassia mimosoides,* Desmodium podocarpum var.japonica,'\ Lespedeza bicolor % and Indigofera pseudotinctoria § asfood plants.Dasyneura viciae (Kieffer) is widely distributed in Europe,including England (e.g. 35), and has been , recorded onnumerous species of Vicia. Full information concerning thisspecies will be found in Vol. II.LETTUCE (Lactuca sativa L.)The only gall midge species that has been found so far on thecultivated lettuce is an undetermined species of Clinodiplosis.* Cassia acutifolia Delile is the source of the best commercial senna drug.t Desmodium gyrans DC is the telegraph plant, and several species are cultivatedfor their use as fodder.J Lespedeza striata Hook. & Am. is a fodder plant.§ Other species furnish indigo.

60 Gall Midges of Economic ImportanceLarvae of this gall midge were found during October, 1939,among lettuce seeds at Slough, England, by J. A. Reid. Thereare specimens in the author's collection. No adults have yetbeen reared. The exact part these larvae play in the economyof the plant is unknown, ye£ it is safe to say they are probablynot injurious.'However, several gall midges have been found on wildlettuces in Europe and the United States of America, includingone potential seed pest of cultivated lettuce. On L. virosa L.(Acrid Lettuce) Rudow (179) recorded Cecidomyia podagrae Bs,which caused a swollen grey mass on this plant in Germany.I have been unable to trace any other reference tothis species.A more important discovery has been made by E. Noury inFrance. He has reared adults of an undetermined Contariniaspecies from larvae found in the capitulum or flower-head ofL. virosa at Tosny. The author has on loan in his collection slidesof both males and females which emerged during October,1936, and on May 26, 1937. If this species ever changes its foodplant to L. sativa, it may becorhe an important seed pest.Reid, who found the Clinodiplosis mentioned above, could findno trace of the presence of a Contarinia in the soil under thelettuces although he made a special search. He was asked todo this as many Clinodiplosis species live as inquilines, and itwas thought possible that the Contarinia might have beenpresent and descended to the soil by the time he found theClinodiplosis larvae.In the U.S.A., Felt (55) described Lasioptera lactucae, whichM. T. Thompson had reared from a tumor gall on the top ofwild lettuce stalks in Massachusetts.MUSHROOM {Psaliota {Agaricus) campestris Quel, orP. (A.) arvensis Quel.)Although several species of gall midges have been associatedwith cultivated mushrooms, few facts are known concerningtheir bionomics. In England it' is generally accepted thatunsatisfactory crops are often-produced on beds where gall

Gail Midges of Economic Importance 61midge larvae are present in large numbers. In AmericaThomas (211) stated that gall midge larvae feed on myceliumin the manure, destroy the bases of mushroom stems andburrow just under the mushroom skin and up into the veil andgills. These parts become slimy, streaked, shredded andyellowish in colour as the result. Thomas adds that canneriesrefuse to accept such mushrooms as the larvae float out intothe broth when the mushrooms are cooked. •Five species have been recorded in England, viz. Mycophilaspeyeri (Barnes), M. barnesi Edwards, ' Pezomyia vanderwulpi(Meijere), Miastor sp. and Lestremia cinerea Macquart, and one,Mycophila fungicola Felt, in the U.S.A.M. speyeri (PI. 7, Figs. 1-2; PI. 8, Figs. 1-2) was originallydescribed (10) in the genus Pezomyia, but was later (13) transferredto the genus Mycophila after it had been compared withthe type of M. fungicola Felt. The larvae were reported by E. R.Speyer as feeding on the mycelium of mushrooms under glass atCheshunt, Herts, England. This latter investigator gave somenotes regarding this midge's behaviour (185). The adults arepositively heliotropic, flying towards the light providing theatmosphere is moist. The greatest number of adults wasobserved towards the end of February. The larva is paedogenetic(18), and the ones that are deiveloping into pupae areorange and possess a black trilobed breastbone. The pupae arenaked and capable of considerable movement. Broekhuizen (34)in Holland records once receiving casing soil containing a greatnumber of orange larvae which he thought were this speciesand stated that the grower could control them by spraying thebeds with salt water. The types and paratypes of the male andfemale as well as a larva with a breastbone from the typelocality are in the writer's collection. In addition, the writerpossesses specimens of females found by P. Bovien (and receivedfrom Matthias Thomsen) in decaying mushrooms at Gentofte,near Copenhagen, Denmark.M. barnesi was described (54) on a male and females caughton windows at Letchworth, Herts. Other examples in the possessionof the British Museum (Nat. Hist.) at the time this specieswas described include specimens caught on windows at Oxford,

62 Gall Midges of Economic Importanceothers reared from mushrooms in April, 1931, at St. Germans,Cornwall (E. R. Speyer), and others reared' from manure atLongsight, Manchester, July, 1932 (M. Shields). Specimensof a male and females reared from manure during August, 1932,at Longsight, Manchester (M. Shields) are in the author'scollection. These are presumably part of the same rearing asthose in the British Museum. The author also has slides oflarvae (including one with a breastbone) and females frommanure in mushroom beds at East Grinstead, Sussex, 1933-34.This species is also paedogenetic.M.fungicola was first described on a male only (59), but laterthe same authority (62) described the female and larva. Theadults were reared in 1897 from young mushrooms in California,U.S.A.This and other related species have been studied by Thomas(2n) who reported'that considerable injury to mushrooms insouth-eastern Pennsylvania, in Ohio, Indiana and other midwesternstates, had been caused by these midges. This species isalso paedogenetic and as many as twenty "daughter" larvaehave been observed within a single "mother" larva.When infested beds are watered thousands of the larvaecome to the surface and either crawl about the soil or congregatein squirming masses often half an inch in diameter. Theyare positively phototropic. Thomas states that, when the soil becomesdry, they have the peculiar habit of coiling and jumping.It is not thought that this method of jumping can be the sameas that of the Contarinia larvae which possess a breastbone(e.g. C. nasturtii, C. lycopersici, etc.). M. fungicola larvae areattracted to moisture and can withstand total submergence inwater for several days (cf. C. nasturtii, p. 42). Drought on theother hand is quickly lethal.The orange and black midges are easily killed by pyrethrumor other dusts and are also strongly attracted to light eitherto traps or to windows. Comparatively few of the larvae reachthe adult midge stage. Picking and destroying infected mushrooms,squashing the larval masses, dusting or spraying withthe usual fly dusts and sprays, e.g. pyrethrum, and drying thesoil surface either with a torch, fan or ventilator or with a

Gall Midges of Economic Importance 63drying agent such as clay or lime are among Thomas'srecommendations for the control of this pest. Hot waterapplied on the surface of the bed has also been successful onoccasion.Thomas reported that in the laboratory several cultures ofthe larvae were killed by an unidentified nematode.Finally, he stated that the larvae on occasion have probablybeen brought into the mushroom houses with the soil or withchopped corn (? maize) stalks when the latter were used inmaking the compost.Pezomyia vanderwulpi (PL 8, Fig. i) was reported by Speyer(i86) to be present with Mycophila speyeri on the mycelium ofmushrooms at Cheshunt. Winged and apterous forms of bothsexes were originally described and figured, as well as the pupaand larva, and placed in the genus Monardia by Meijere (133)from specimens reared from a rotten willow trunk in Holland.Kieffer (107) erected the genus Pezomyia and made vanderwulpithe type species. The author has in his collection specimens offemales of this species caught over a hazel stump near Leedsand, in addition, brachypterous males and a normal wingedfemale reared early in April, 1945, from rotten Spruce Fir woodcollected at Cotterill Clough, Cheshire, by H. Britten. AnAlgerian species, P. brachyptera Kiefiêr, was reared from anunidentified mushroom (107).The undetermined Miastor species was recorded (16, 210) asoccurring in mushroom sheds at West Barnham, West Sussex.Only larvae were found, mostly of the paedogenetic typewithout a breastbone, but some possessed a breastbone whichwas hastiform. In spite of this no adults were reared. The larvaewere pearly-white to pallid yellowish-white in colour. They fedbetween the gills of the "buttons" and maturing mushrooms,sometimes working right up into the cap. The latter whenattacked turned brown and shrivelled. Up to seventy larvaewere taken from the gills of a small mushroom and often tento twenty from the flesh of the cap. The chief damage is doneto the spawn before the beds begin to break. Theobald statedthat these paedogenetic larvae were doing more harm than thelarvae of Sciara agraria which were also present. Specimens of

64 Gall Midges of Economic Importancea larva with a breastbone as well as larvae without are in theauthor's collection. .Lestremia cinerea was originally described by Macquart (125),but the type is lost. Edwards (53) re-described it. It is a commoncentral and north European species and captured specimenshave been recorded from several counties in England—butonly once so far as the author is aware has it been reared.In 1937 E. R. Speyer sent the author specimens which he hadreared in May and June from mushrooms grown out-of-doorsat Eastbourne, Sussex. The larvae were in the stalks and themushrooms were described as being of very abnormal growth.Specimens of males, females and pupae of this rearing are inthe author's collection, as well as a male and females caughtin a mushroom house at Worthing, Sussex, by F. C. Woodduring 1936.PARSNIP {Pastinaca sativa L.)There are three European species that live on parsnip, viz.Contarinia pastinacae (Riibsaamen), Kiefferia pimpinellae (F. Loew),both of which destroy the seeds, and Macrolabis corrugans(F. Loew), the larvae of which live on the leaves.K. pimpinellae has already been dealt with as a pest of carrots(pp. 27-9), C. pastinacae and M. corrugans receive attentionbelow. The latter is particularly interesting since, although itis exceedingly common on a number of wild Umbellifers, onlyvery occasionally has it been reported as attacking cultivatedparsnip. It has been suggested (20) that the rate of unfoldingof the leaves may very well be responsible for this (see belowunder DAMAGE, p. 65).Contarinia pastinacae was originally described by Riibsaamen(166) from Germany. Itis a typical Contarinia, and the larvae areyellow. The latter live in the slightly swollen fruit of parsnip{P. sativa), the seed being consequently destroyed. No otherfood plant has been mentioned in the literature. The onlyreference to this midge in economic papers is that of Noel (146),who gave it in his list of parsnip pests in France.

Gail Midges of Economic importance 65lâcrolabis corrugans (F. Loew, 1877),syn. Cecidomyia heradei Kaltenbach, 1874.DIAGNOSTIC CHARACTER. Leaves of parsnip crinkled, remainingfolded and unopened and later blackening (PL 9).DAMAGE. When badly infested the leaves are completelyfolded up and swollen at the points where the larvae are to befound. The leaves never unfold, become black and finally thetissue rots. Naturally the development of the root is seriouslyaffected. Fortunately this midge has only been reported oncultivated parsnip occasionally, e.g. by Rubsaamen (164) inGermany, Noel (146) in his list of parsnip pests and three timesin Great Britain. Hall (80) recorded it on parsnip in Kent,Wildman (224) reported it from Yorkshire and stated that insome instances the damage was very severe, and the thirdinstance is a severe attack near Edinburgh in 1932.The rarity of this midge's occurrences on cultivated parsnipin comparison with its almost perpetual occurrence onHeracleum sphondylium is noteworthy. The author in 1933-34carried out a series of experiments in which he allowed midgesreared from H. sphondylium to have their choice of ovipositionon this plant and on "Hollow Crown" parsnip. Although themidges oviposited frequently on the unopened leaves of thecultivated plants, in only a few instances did the larvae establishthemselves. On the other hand, the typical malformationseemed to result from every oviposition on the wild plants. Itwas noticed that the rates of unfolding of the leaves on the twospecies of plant were very different; whereas those of H.sphondylium opened out quite slowly, those of the "HollowCrown" unfolded comparatively fast. The eggs and very younglarvae of the midge became exposed to the direct sunlight onthis latter plant, whereas on the wild plant they always hadsufficient time to establish themselves arid begin their feedingactivities which prevented the leaves from unfolding. Thisdifference in rate of unfolding of the leaves, together with theflifference in their superficial texture, seems to afford a logicalreason for the observed difference in frequency of infestationof cultivated parsnip and Heracleum sphondylium.DESCRIPTION. F. Loew (124) originally described this midge,

66 Gall Midges of Economic Importancealthough Kaltenbach (92) had previously given the nameCecidomyia heraclei to what is probably the same gall. Themidge is a typical Macrolabis. The abdomen of the female ispale orange-yellow in newly emerged specimens. The larvaeare white.DISTRIBUTION. Central and western Europe, includingFrance, Holland (136), Germany (119), Italy (214) and GreatBritain. In the last-named country it is widespread throughoutEngland (3) and occurs in Wales, Scotland and Ireland, e.g.in Co. Tyrone and Co. Armagh (126).LIFE HISTORY. The female midge lays her eggs in the foldsof the leaves before they open. In a few days the larvae hatchand as a result of their feeding the leaves do not unfold. Thewhite larvae which are gregarious are fully fed after about amonth. By this time the infested leaves often have becomeblackened. The larvae descend to the soil to pupate. After theyhave left, the leaves, if not completely blackened and rotted,unfold revealing scars where the larvae have fed. Such scarsare a common sight on the older leaves oi Heracleum sphondyliumthroughout the summer from June onwards.In the spring of 1933 midges appeared on the wing fromApril 18 to June 2 at Harpenden, Hertfordshire. Some of the «first generation larvae resulting from the midges were fully fedand had descended to the soil for pupation by May 28. Theadult midges of this generation emerged between June 13 and19. Second-generation midges were on the wing from July 26to August 17. Some third-generation larvae were fully grownby September 3 and the adult midges of this generation emergedbetween May 10 and 24, 1934. Thus there are three generationsin some years at least.FOOD PLANTS. In addition to Parsnip (Pastinaca saliva L.),several Umbellifers, including Common Cow-parsnip or Hogweed{Heracleum sphondylium L.) and the Chervil {Chaerophyllumaromaticum L.), from which the species was originally described.Other host plants in this family are the Hog's Fennels {Peucedanumcervaria Lapeyr. and P. orseoselinum Moench) and possiblyChaerophyllum hirsutum yUl. (4.2). rThere are other food plants'mentioned in the literature, but

Gall Midges of Economic Importance 67as they do not belong to the UmbeUiferae it would be wiserfor the present to regard them with a certain amount of doubt.It is most unusual for a single gall midge species to occur onplants belonging to more than one family.Originally, Riibsaamen (164) recorded M. corrugans onWhite Dead-Nettie {Lamium album L.). Later (165) he statedthat he could not separate the midges causing terminal budmalformations on Red Dead-Nettie [L.purpureum L.) and WhiteDead-Nettie from M. corrugans and so referred to them asM. corrugans. In 1915 (175), however, he was able to see differencesand described the species making the terminal bud galloi Lamium spp. as M. lamii. Trotter (215) also recorded M.corrugans on another Dead-Nettie {L. flexuosum Ten.), whileHouard (85) considered that the midge which Cotte in 1912had recorded from a Dead-Nettie, L. inaculatum L. var. rugosumBriquet, might be M. corrugans. It is most probable that theseauthorities were in reality recording M. lamii Rubsaamen (175).Likewise, Swanton (191) recorded M. corrugans, referring toConnold (41), on Savoy Hawkweed {Hieracium boreale Fries),whereas he should probably have recorded M. hieracii KieflFer(96, 104). Bagnall & Harrison (3) recorded M. corrugans onHedge Woundwort {Stachjs sylvatica), but it is considered mostunlikely that this is a food plant of the true corrugans.NATURAL ENEMIES. Anthocoris nemorum L. has been noticedfrequently sucking the larvae on the leaves of Hogweed.An unidentified Chalcid parasite has also been reared by theauthor.CONTROL MEASURES. It has been suggested (20) that theapplication of suitable artificial manures to promote rapidgrowth might be beneficial in thwarting an attack.MATERIAL. Males and females, England (Yorkshire, onP. sativa) and Wales (Caernarvonshire, on H. sphondylium).Larvae, Scotland (Edinburgh, on P. sativa).MOST IMPORTANTREFERENCESLoEW, F. 1877. Verh. zool.-bot. Ges. Wien, Tt, 11-13 (Cecidomyia)BARNES, H. F. 1934. Sci. Progr., 29, 84.

68 Gall Midges of Economic importancePEA {Pisum sativum L. and P. arvense L.)Only one gall midge has been reported as a pest of peas.This is Contarinia pisi (Winnertz), which is a European form.It has been a major pest, in the' pea-growing areas of severalcountries, especially in connection with the pea-canningindustry of Switzerland. From the published records it appearspossible that peaks of abundance are apt to occur about everyfifth year. There are reports of serious damage being done bythis midge in various countries during 1905, 1915, 1925 and1926 (two countries), in 1931 and in 1938 and 1939.Contarinia pisi (Winnertz, 1854), the Pea Midge,*syn. piskola Meijere, 1911.DIAGNOSTIC CHARACTER. Swollen flowers, flower stemsremaining very short and clustered together and slightlymalformed pods of peas and perhaps beans, in which are to befound white "jumping" larvae.DAMAGE, There are thus three points of attack, viz. theflowers (PI. 10, Fig. i), the terminal shoots (PI. 10, Fig. 2)and the pods (PI. 6, Fig. 2). In infested flowers, the basalparts, especially of the sepals are swollen, the petals are verycrinkled and sometimes scarcely visible at all, and the stalks maybe of normal length, as illustrated by Barnes (12) and Kutter& Winterhalter (114). Up to ninety-nine larvae have beenfound in such a single swollen flower. When the terminal shootsare attacked, the flower stems remain about half an inch inlength arid clustered together; this type of damage is alsoillustrated in the above-mentioned papers. In parts of Kent it isknown as "Nettlehead".Both these types are much more serious than the third, theinfested blossom usually dying vvithout forming pods at all.However, sometimes small, dwarfed and malformed pods areformed, but the peas do not develop.Serious losses due to these two types of injury have occurredin the Canton of St. Gall, Switzerland, to the peas grown forcanning. Some idea of the numbers of midges present can be* Other popular names include the following: Hernesaaski, Aertgallmyggen,Erbsengallmiicke. "

Gail Midges of Economic importance 69gained from the statement that up to 126 cocoons were foundper square yard, which is roughly 620,000 per acre, or the lossof 20,000 pods. In fact, these types seem to be the better knownin Holland and Switzerland; in the last-named country the podtype of damage was not encountered (i 14). In England, on theother hand, until 1927 C.pisi had not been generally recognizedas responsible for the swollen flowers and clustered terminalshoots, although such injury is common and widespread andHarrison (83) had mentioned the fact.*When the pods are attacked, usually 20 to 40 larvae arefound in a single pod, although as many as 300 have beenrecorded (206). Infested pods sometimes have a gouty orswollen appearance. The larvae seem to feed on the innersurface of the pods and even on the peas themselves. Thedamage is then more of a nuisance than of real importance,the peas rarely suffering noticeably. It is nevertheless unpleasantto find the larvae among the peas when served at table and alsoamong shelled peas for market; in both instances their sale priceis consequently lessened. Smith (184) had a good illustrationof this type of attack, which is reproduced in PI. 6, Fig. 2.DESCRIPTION. Although Malpighi and Reamur are stated tohave referred to this midge in 1687 and 1737 respectively,• Winnertz (226) was the first to name it. Both sexes of thistypical dirty-coloured Contarinia, which is of average size forthis genus, and the milky-white larvae were described.SYNONYMY. Meijere (135) described C. pisicola, separating itlargely on account of the supposedly different habit of attackingthe terminal shoots. Rubsaamen & Hedicke (178) retainedpisicola Meijere separate from pisi Winnertz. The author hasbeen honoured by the gifts of some of Meijere's specimensobtained in 1909-10 and also specimens obtained by Kutter inhis investigations. After a comparison of these Dutch and Swissspecimens with English specimens, there remains no doubt butthat C. pisicola Meijere and C. pisi Winn, are one and the samespecies. The name given by Winnertz has priority.DISTRIBUTION. Widespread in Europe, having been recorded* Owing to misreading, the author (12) stated that Harrison had not mentionedthe species of Contarinia involved, but this was a mistake.

70 Gall Midges of Economic Importancefrom France (102), Italy, Switzerland (114), Holland (136),Germany (226), Norway (183), Sweden (219), Denmark (i2i),Finland (74) and England and Wales. In England and Walesit probably occurs in every county,' being recorded in Englandfrom Kent and Worcestershire (206), Northumberland andNorfolk (7), and having been found in Cornwall, the Isle ofWight, Hampshire, Surrey, Middlesex, Berkshire, Cambridgeshire,Lincolnshire, Leicestershire, Derbyshire, Nottinghamshire,Lancashire and Yorkshire. In Wales it has been foundin Caernarvonshire.LIFE HISTORY. The bionomics of this midge 'in Switzerlandhave been studied in detail by Kutter & Winterhalter (114)and Kutter (i 11, 112, 113). Theobald (206) and the author (13)and Meijere (135) have made shorter studies of it in Englandand Holland,The eggs are laid in batches of twenty or more inside theflowers, on the young leaves or shoots, and perhaps in the pods,although the exact manner in which the larvae get inside thepods is unknown. The larvae hatch after about four days. Ifthe flowers are attacked, they live at the base of the ovaries;in terminal shoots attack they are to be found among theclustered leaves; and where the pods are infested the larvaelive inside the pods.The minimum duration of the larval stage is about ten daysfor most of the first generation,* while some of this generationand all of the second do not pupate until the following spring.When fully fed the larvae "jump" to the soil where they spincocoons in which they pupate. These cocoons can be founddown to a depth of 3! in. The pupal stage lasts about elevendays and Kutter (iii) states that adults live six days, whichif it is the rule, is rather a long period for adult midges.Taschenberg's statement that the adult midges live throughoutthe winter cannot be substantiated.Both in England and Switzerland there are two generationsa year, but probably the first flight of the year includes someindividuals of the previous year's first generation in additionto those of the second. In England the adults of the second* A generation is considered to start with the egg and end with the adult midge.

Gall Midges of Economic Importance 71generation of 1925 (or the first flight of 1926) were on the wingduring the last two weeks of June in 1926, while those of thefirst generation of 1926 (or second flight of 1926) were flyingduring the last two weeks of July and the first week in August.In Switzerland the flight periods are slightly earlier, startingat the end of May in some years, although the midge's firstappearance may vary a fortnight from year to year.Kutter (112) stated that during 1934, in contrast to allprevious experience and expectations, all the overwinteringlarvae did not emerge as adults at the end of May or beginningof June. In fact, by far the greatest number emerged a fortnightlater in the second half of June and up to mid-July. He suggeststhat this binodal curve of emergence may be due to some of thefirst generation of the previous year aestivating and carryingover to the next year when they emerged earlier than themidges derived from the second generation of the year. Therewere thus three flights of midges in 1934.The same phenomenon of the first flight of the year havingtwo crests has been observed by the author for one of the WheatBlossom Midges [Sitodiplosis mosellana Gehin). In 1931, 1933 and1939 the peaks were respectively June 14 and 28, June 5-6and July 3, and June 15 and July 7.Incidentally it has been definitely proved that some individualsof both the Wheat Blossom Midges (S. mosellana andC. tritici Kirby) sometimes pass more than one winter in thesoil before emerging (22). There is, however, no proof yet thatthe older midges emerge earlier in the season of emergence.Certain species, such as the Pear Leaf Midge (Z). pyri Bouche)and the Arabis Midge {D. alpestris Kieffer), have severalgenerations a year, and are known to have partially aestivatingand hibernating generations. In no instance, however, do theauthor's daily figures of emergence show any indication ofa binodal curve, or of the midges of the earlier generationsemerging earlier. In these instances the larvae of the earliergenerations were kept separate.The Button-top Midge of Willows {R. heterobia H. Loew) isalso known to have at least two generations a year, and theweekly emergence figures for 1936 and 1938 show a binodal

72 Gall Midges of Economic Importancetendency, but an examination of the daily figures of theseparate samples. do not support this view. It is, however,extremely unlikely, owing to the method of collecting, thatmany galls of earlier generations appeared in the samples.Kutter (lis) also suggested that cultivation might be partlyresponsible for the double emergence; ploughing would burythe cocoons deep and so the midges would be affected by thespring weather later than those nearer the surface, while theemerging midges would have further to come on their passageout of the soil (see section on NATURAL ENEMIES, p. 73).FLUCTUATIONS IN NUMBERS. It is possible that Contarinia pisihas a somewhat regular rhythm in abundance, noticeable outbreaksin England having occurred in 1905 and 1906 (adS),and in 1926 (222); in Sweden in 1915 (219); in Holland in1910 (135) and 1925 and 1926 (91); in Norway in 1924 and1925 (183), and 1938 and 1939 (90); and in Switzerland in1931 and 1932 (114).FOOD PLANTS. Green Pea {Pisum sativum L.) and Field Pea(P. arvense L.). It may also occur on beans, since A. W. RymerRoberts {in lift., 2.viii.i929, 3.12.40) has found a Contariniaspecies in the pods of broad bean [Vicia faba L.). It is alsopossible that it will be found on Sweet Pea {Lathyrm odordtus L.),since Clinodiplosis pisicola Barnes, its inquiline midge, wasoriginally described from this plant.RESISTANT VARIETIES. From time to time it has been statedthat one class or other of peas is more likely to be attackedthan another. For example, it has been stated that early peasdo not seem to be attacked, whereas main crops suffer most(206). Harrison (83) stated that late varieties like Gladstonewere unharmed, and earlies and second earlies such as Senatorand Leader were heavily infested. The author (12), writing ofthe first generation, stated that of peas examined on July 6,1926, the earlier sowings suffered most, those varieties in fullflower about the first week in July actually being most heavilyattacked. Those that had not flowered by this date were themost free, but those that had already podded were found tobe infested to a slight extent in the pods.,Itseems probable that all varieties of peas are subject to attack

Gall Midges of Economic Importance 73and that there is no real varietal resistance. What immunitythere appears to be is probably caused by an isolation in timebetween the plant variety and insect. The extent to which anyparticular variety or class of pea will be attacked depends onwhether the plant is at one of the stages of growth suitable foroviposition and successful infestation at the time when theadult midges are abundant. This will vary in different partsof a country, and exceptionally from season to season. Unlesstherefore one has a long experience of dates of emergence ofthe midges, and an extensive knowledge of the phenology ofthe different varieties of plant under consideration, it would bevery unwise to attempt to dogmatize on which varieties arelikely to be more attacked than others.NATURAL ENEMIES. Four parasites have been recorded,namely the Scelionids, Sactogaster pisi Forst., Inostemma bosciiJur. and Leptacis tipulae Kirby, and the Pteromalid, Pirenegraminea Hal. Kieffer (109) redescribed S.pisi, while Kutter &Winterhalter (114) found all four species in Switzerland, onlyS. pisi and P. graminea, however, being common. Details of thebionomics as well as illustrations of the parasites are to be foundin the last-cited paper as well as in those of Kutter (m, 112,113). Husas (90) only found S. pisi in southern Norway.S. pisi oviposits in the eggs of C. pisi and its larvae were foundin the larvae of the latter in their cocoons up to the end ofOctober. In 1932 up to 50 per cent, of the midge's larvae wereparasitized. S.pisi was abundant again in 1933 during July, andagain more so in 1935, when up to 50 per cent, of the secondsummer generation of C. pisi was attacked.A detailed account of the morphology and biology of P.graminea is given by Kutter (i 11). It has two generations a yearfollowing those of C. pisi which it resembles by having a binodalfirst-emergence curve. This may be due to some of the firstgeneration carrying over to the next year and/or to the effect ofploughing. In one instance P. graminea emerged 8 days earlyfrom unploughed land as compared with some from ploughedland. The eggs are laid in the larvae of the midge. The primarylarval stage lives on the material in its tgg, while the secondstage develops at the expense of the midge larva. The full-

74 Gall Midges of Economic Importancegrown larva leaves that of its host and pupates in the latter'scocoon. The second and some of the first generation overwinterin the primary larval stage. The minimum period required bythe parasite to complete a generation in the summer at roomtemperature was found to be 24 days. No hyperparasite wasfound. P. graminea was considered by Kutter to be responsiblefor the comparative scarcity of the pea midge in 1935, theextent of parasitism of this species having been 50 per cent, in1932, 75 per cent, in 1933 and 85-88 per cent, in 1934. In 1935this parasite decreased for lack of hosts./. boscii was commoner in 1933 than in 1932, but L. tipulaewas not seen in 1933.[Smith (184) states that the Scelonid Polymecus hopkinsi Crawf.& Brad, has been recorded as a parasite of the pea midge inthe United States of America. This record must be regardedwith grave suspicion, since the author cannot verify the existenceof the pea midge in the U!S.A.]In addition to the above-mentioned parasites, one predatorhas been recorded, namely the gall midge Lestodiplosis pisiBarnes (17). The larvae of this midge are bright red and feedon those of C. pisi. They were found in Kent.iNquiLiNES. The larvae of the gall midge Clinodiplosis pisicolaBarnes* live as inquilines in the pods of the garden pea infestedby Coniarinia pisi. This species was originally described (15)from larvae found in the pods of sweet pea [Lathyrus odoratus L.)at Aberystwyth, Wales. It has beenfound in addition in companywith larvae of C. pisi in pods of garden peas at Harpenden,Hertfordshire and at Barnstaple, Devon.CONTROL. The use of insecticides against this midge are notsatisfactory. Spraying with nicotine and soap (112) was ineffective,and in addition, killed large numbers of the beneficialparasite Pirene graminea. Likewise Husas (go) found thatspraying with nicotine sulphate (i : 400 or i : 600) was not veryeffective. Various workers have suggested the use of naphthalene,e.g. Barnes (12), at the rate of 3 cwt. per acre, against the larvaein the soil, but no good results have been recorded from this* This species may later prove to be„synonyiiious with Clinodiplosis lathyriRiibsaamen (176),

Gall Midges of Economic Importance 75treatment. Similarly, Kutter (112) found the application ofkainit unsatisfactory.Early sowing and the selection of early varieties are claimedby Husas (90) to be the best preventive methods (but seeRESISTANT VARIETIES, p. 72).Cultural methods of control, on the other hand, have proveduseful. Theobald (206) recommended burning all infestedplants as soon as the crop has been gathered. This destroys anylarvae left in deformed pods and shoots. He also suggested thatthe land should be deeply trenched, rolled and only top-workedthe next spring. This latter was presumably to ensure that thecocoons remained deeply buried.Similarly, Kutter (112) found that ploughing either in theautumn or in spring reduced infestation. Thus, the midge is afurther example of a gall midge pest being controlled by cultivationat the correct time of year in relation to its life history.It would be reasonable to suppose that intensive and nicelytimed cultivation during the summer would so disturb thefirst-generation full-grown larvae that they could not developsuccessfully, and in this way the second generation would beprevented.Crop rotation is also satisfactory. Kutter & Winterhalter(114) stated that in Switzerland crop rotation had been madecompulsory, with the result that areas under peas in 1931 and1932, but not in 1933, were at first entirely free from midgewhen peas were again grown in 1934, and later were onlypresent in small numbers.Husas (90) recommended deep digging in the autumn, theuse of new land at a distance from old areas, and only growingpeas once in four years.In small gardens, however, handpicking the infested shoots,flowers and pods would probably be more effective than rotationof the rows of peas.Natural barriers, such as woods and hills and even an embankment15 ft. high, were found by Kutter (m) to beefficient in preventing the spread of the midge. He suggestedthat in open country peas should not be grown within i kilometreof infested areas.

76 Gall Midges of Economic ImportanceKutter (in) also found that trap-crops of peas could be usedto intercept flying midges. IBIOLOGICAL CONTROL. Kutter (11,2) suggested that the introductionof the Pteromalid Pirene graminea into areas infestedby the pea midge might prove beneficial.MATERIAL. Males, females and larvae, England (Cambridgeshire,Devon, Hertfordshire, Kent), Switzerland (from H.Kutter) and Holland (from J. C. H. de Meijere, specimensobtained in 1909-10 and identified by him as C. pisicolaMeijere). Pupa, Holland (from Meijere). Parasite, Pirenegraminea Hal., Switzerland (from Kutter). Predator, Lestodiplosispisi Barnes, England (Kent) (males, type and the 2 para types;females, type and the 4 paratypes). Inquiline, Clinodiplosispisicola Barnes, England (males, Devon—females, Devon andHertfordshire—larvae, Hertfordshire), Wales (Cardiganshire)(type and the 8 paratype males; type and the 9 paratypefemales; pupae; larvae).MOST IMPORTANTREFERENCESWiNNERTZ, J. 1854. Stett. Ent. Zît., IS, 325-7 (Cecidomyia pisi)MEIJERE, J. C. H. DE. 191 I. Tijdschr. Ent., 54, 183-7 {Contannia pisicola)THEOBALD, F. V. 1907. J.S.-E. agric. Coll., Wye, no. 16, 135-8 {Diplosispisi)BARNES, H. F. 1927. J. Minist. Agric., 34, 159-61 (Contarinia pisi)KUTTER, H. & WINTERHALTER, W. 1933. Landw. Jb. Schweiz-, 47, 273-338{Contarinia pisi)KUTTER, H. 1934. Mitt. Schweiz. ent. Ges., 16, 1-82KUTTER, H. 1934. Landw. Jb. Schweiz-, 48, 1133-72KUTTER, H. 1936. Landw. Jb. Schweiz-, 50, 80-102. .POTATO iSolanum tuberosum L.)The only gall midge recorded from potato is Asphondyliatrabuti Marchal, whose larvae live gregariously in the fruit, nottubers. Marchal (129) received living galls from Algiers at theend of September just when emergence was taking place.Consequently he was able to watch this and describe the femaleand the pupa. He did not see the larva.

Gall Midges of Economic importance 77RHUBARB {Rheum rhaponticum L.)The larvae of an unidentified gall midge live in slightlyraised cells on the stalks of rhubarb. A single larva live? in eachcell. The only possible damage this species can do is to make therhubarb unsightly and so lower their market value slightly. Sofar this midge has only been reported from Newry, NorthernIreland, but it is very probable that it is much more widelydistributed. It may be a species of Profeltiella. The author hasa single specimen of a larva received from R. G. Chamberlain,who is responsible for its discovery in 1937.SQUASH {Cucurbita pepo L. var.*)The male and female of Cecidomyia cucurbitae (Felt) weredescribed by Felt (58), who recorded that they were rearedfrom orange larvae from a squash having a curious roughfulvous appearance. Later the same authority (68) mentionedthat this species was reared in July, 1876, presumably by C. V.Riley but gave no locality. Felt also stated that Mycodiplosiscucurbitae Felt (58) was associated.SWEET POTATO {Ipomoea batatas Lam.)One species of gall midge has been recorded from thisvegetable, namely Schizomyia ipomoeae Felt (56). Both sexes andthe empty pupal skin, and later (57) the larva, were described.It was originally reared by W. H. Patterson of St. Vincent,B.W.I., in large numbers from the flower-buds. S. ipomoeae isunusually small for a species of this genus, being only i'2 to1-75 mm. in length. The same information is given again byFelt (63).Rtibsaamen (171) had previously described a leaf gall containingyellowish larvae belonging to the genus Clinodiplosis onIpomoea sp. from Brazil.* Cucurbita pepo L. is the pumpkin, with its varieties the vegetable marrow andsquash. C. maxima Duchesne, the giant pumpkin, is also cultivated in NorthAmerica.

78 Gail Midges of Economic ImportanceTOMATO (Solanum lycopersicum L.)Only one gall midge attacks tomato, but this one, Contarinialycopersici Felt, is a local major pest. Fortunately for growers,this midge is so far restricted to the West Indies, British Guiana,British Honduras and possibly Hawaii.C. solani Riibsaamen has also been recorded (192I fromtomato flower-buds in Hawaii but probably in error for C. lycopersiciFelt (21).Contarinia lycopersici Felt, 1911, the Tomato Flower Midge.DIAGNOSTIC CHARACTER. Yellowish-white larvae in unopenedflower buds of Tomato.DAMAGE. The flower-buds remain closed and usually dropprematurely. Sometimes the ovary is damaged, resulting inmalformed fruit. But Topper (213), after inoculating unopenedbuds with larvae of this midge, came to the conclusionthat they were not responsible for "early" damage to thetomato fruit. This midge is not generally distributed in theWest Indies, but where it occurs it is considered a majorpest (36). •DESCRIPTION. Felt (60) first described this species. It is arather small Contarinia species, the females being only aboutI mm. in length. The abdomen is brownish. The larvae areyellowish-white.DISTRIBUTION. This midge was originally described from St.Vincent (60) and has since been recorded from Trinidad (19),Barbados and Hawaii (21), Grenada (36) and St. Lucia,Dominica, British Guiana and British Honduras (37).LIFE HISTORY. Brief descriptions of the life history were givenby Patterson (149) and Briant (33), but the fullest accoimt isthat by Callan (37). The female midges lay their eggs in theearly evening (5.30-6.45 p.m.) in young tomato flower-budswhen the latter are only about i-2 mm. in length. The larvaeprobably hatch in about 2 days and feed on the anthers, styleand ovary, Patterson recorded, 5-6 or more larvae in a singleflower; Briant said 8-10; while Callan found as many as 60—

Gall Midges of Economic Importance 79but the average per sample varied from slightly over 2 to i6,giving a total mean of about 7. In one sample as many as83 per cent, of the flowers were infested.At first the larvae are white but when full grown they arebright yellow. They "jump" out of the flower-buds to pupateabout I in. below the surface of the soil. The larval stageprobably lasts 8-10 days, while the pupal stage is 9-10 andoccasionally 12 days. The total life cycle is about 18-22 days.The larvae are very susceptible to humidities much belowatmospheric saturation, and successful pupation demands dampsoil.Callan (37) records finding larvae during June, July, August,September, November and December. Few larvae were in thetomato blossom during the dry season (when tomatoes areusually grown in Trinidad), but in the wet season nearly everyflower-bud may be infested.FOOD PLANTS. The normal host plant is the Tomato {Solanumlycopersicum L.), but Holdaway (84) cites this species as a submajorpest in Hawaii of Egg-plant and Pepper as well as ofTomato. Further evidence of the correct identification of themidge species involved on these plants is desirable.*PARASITES. A Sactogaster sp. has been found in Trinidad byCallan (37), who also obtained what appeared to be identicalfrom Grenada.CONTROL. Cultural methods of control, based on the requirementof damp soil by the larvae for successful pupation, arerecommended by Callan (37). He states that the crop shouldbe planted so as to flower only in the dry season, the soil roundeach plant should be kept well forked and watering should belimited to the space between the rows of plants so as to wet thesoil around the base of the plants.Callan (37) stated that it had been reported that Vaporiteapplied to the surface of the soil appeared to give some control,while he suggested tentatively the use of naphthalene andparadichlorobenzene. He also suggested spraying with nicotine* It would appear from some very recent and hitherto unpublished studiesby D. D. Jensen of the University of Hawaii Agricultural Experiment Station thatonly one species is involved.—i Nov., 1945.

80 Gall Midges of Economic Importanceand pyrethrum in the early evening when the midges areovipositing.ICLOSELY ALLIED SPECIES. The European Contarinia solaniRiibsaamen (i66) whose larvae prevent the flower-buds ofWoody Nightshade {Solarium dulcamara L.) from opening.Swezey (192) reported this midge from Hawaii, but probablyin error for C. lycopersici (21). Specimens of males and females(Kent) and larvae (Hertfordshire) of C solani are in the author'scollection.MATERIAL. Males and females, Trinidad and Barbados.Larvae, Barbados. (All from Tomato.)MOST IMPORTANTREFERENCESFELT, E. P. 1911. Ent. News, 22, 303-4CALLAff, E. McC. 1941. Trap. Agriculture, Trin., 18, irg-25.VEGETABLE MARROW {Cucurbita pepo L. van*)In Europe Jaapiella bryoniae (Bouche) has once been reportedas a pest of vegetable marrow. This is a widely distributedEuropean gall midge that usually infests the terminal leavesand growing points of White Bryony {Bryonia dioica Jacq.).Perhaps the reason why it is not found more frequently onvegetable marrow is that it is the normal custom to nip out thegrowing points quite early in the season.Jaapiella bryoniae (Bouche, 1847).fDIAGNOSTIC CHARACTER. Terminal growing point clusteredand malformed, developing into a rosette with a hard core.DAMAGE. The growing point is destroyed. The only recordof this midge as a pest of vegetable marrow and pumpkins isthat of Noel (145) in France.DESCRIPTION. Bouche (31) originally described this speciesfrom Bryonia alba L. The abdomens of the females are palered when newly emerged; otherwise the general appearance ofboth sexes is brownish.* See footnote, p. 77.t This midge has been called the. White Bryony and Vegetable MarrowMidge (11).

Gall Midges of Economic Importance 81DISTRIBUTION. Central and western Europe, includingPortugal (195), France,(118), Holland (136), Germany (92),Italy (215), Hungary (193) and Great Britain. In the lastnamedcountry it has been found in Hertfordshire near Letchworthby Bagnall & Harrison (4), and at Harpenden by theauthor. It has also been recorded from Algeria (85).LIFE HISTORY. The females lay their eggs in the terminalshoots among the developing leaves. The larvae, which arewhite and gregarious, feed in the galls, which soon develop.When fully fed, presumably in about a month, they leave thegalls and pupate in small whitish cocoons in the soil. Pupation,which in the summer takes 8 to 14 days (31), may be delayedtill the spriifg in the overwintering generation. It seems likelythat the overwintering generation appears on the wing in June.There is probably another flight in July, and in some yearsthere is a, further flight in August and September. In 1940 theauthor found- full-grown larvae, as well as other larvae ofvarying size, on August 12. On August 30 midges started emergingand continued until September 6, after which .there wereoccasionally emergences until October 11. In 1941 emergencestarted again on May 29 and continued until June 21. Gallson White Bryony were readily visible on June 17, 13 daysafter oviposition on June 4. By June 21 the larvae were abouthalf-grown. There are probably two or three overlappinggenerations a year.FOOD PLANTS. White Bryony [Bryonia dioica Jacq.) andB. alba L. It has been recorded once from Vegetable Marrow(145)-The author has carried out preliminary trials with specimensreared from B. dioica to see whether they will attack vegetablemarrow. At first no interest was shown by females when theywere placed in the presence of vegetable marrow shoots alone.Later a shoot of White Bryony was inserted in the cage and allthe female midges became excited,and began to oviposit on itand the vegetable marrow shoots. Although the bryony shoothad been extracted overnight the midges still continued ovipositingindustriously throughout the next day on the vegetablemarrow shoots.

82 Gall Midges of Economic ImportanceNATURAL ENEMIES. Although none has been recorded, severalunidentified Chalcid parasites have been reared from larvaefound at Harpenden.CONTROL. Picking and destroying the infested shoots as soonas they are recognizable would be effective in reducing attacks.MATERIAL. Males, females and larvae from B. dioica,Harpenden, Hertfordshire.MOST IMPORTANTREFERENCEBoucHE, P. FR. 1847. Stettin, ent. Ztg-> 8, 144 {Cecidomyia).YAM {Dioscorea spp.)•Riibsaamen (172) recorded small leaf galls, which areequally visible on either surface of tlie leaf, on Dioscorea minutifoliaEngl, in the Cameroons. He figured the larval breastbone.The author has in his collection male specimens of a Trifilamidge and female specimens of a Lasiopterine midge reared byE. Hargreaves from leaf galls on Yam at Njala, Sierra Leone,during 1936.In addition, Felt (65) figures an irregular warty stem gallon Yam which he attributes to an unidentified Cecidomyiaspecies. He gives no other information.

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92 Gall Midges of Economic Importance200. TAVARES, J. S. 1920. "Cecidologia Brazileira. Cecidias que se criamem plantas das familias das Leguminosae, Sapotaceae, Lauraceae,Myrtaceae, Punicaceae, Aurantiaceae, Malpighiaceae, Sapindaceae,Umbelliferae, Loranthaceae, Apocynaceae, Urticaceae, Salicaceaee Gramineae", Broteria, zool., 18, 82-125.201. 1922. "Cecidologia Brazileira:' As restantes familias", Broteria,zool., 20, 5-48.202. 1924. "Noticias Cecidologicas",'Bro/erw, zooL, 21, 91-8.203. 1925. "Novas contribuigao para o conhecimento da CecidologiaBrazileira", Broteria, zool., 22, 5-55.204. TAYLOR, T. H. 191a. "Cabbage-top in Swedes", Univ. Leeds & Tork.Counc. agric. Educ, no. 82, 21 pp.205. THEOBALD, F. V. 1892. "An Account of British Flies (Diptera),Vol. I", Elliot Stock, London, xix + 215 pp.206. 1907. "Report on Economic Zoology for the year ending April1st, 1907", J.S.-E. agric. Coll. Wye, no. 16, 29-180.207. 1910. "Report on Economic Zoology for the year ending September30th, 1910", J.S.-E. agric. Coll. Wye, no. 19, 73-211.208. 191 !• "Report on Economic Zoology for the year ending September30th, 1911", J.S.-E. agric. Coll. Wye, no. 20, 89-244.209. 1912. "Report on Economic Zoology for the year ending April1st, 1912", J.S.-E. agric. Coll. Wye, no. 21, 111-220.210. 1928. "Notes on a new Mushroom Pest and on the control ofSciara larvae", J.S.-E. agric. Coll. Wye, no. 25, 79-81.211. THOMAS, C. A. 1942. "Mushroom Insects: their Biology and Control",Bull. Pa. agric. Exp. Sta., no. 419, 43 pp.212. THOMAS, D. C. 1946. "A Study of the distribution of the SwedeMidge (Contarinia nasturtii Kieffer) in Devon and symptoms of itsattack on various Plants", Ann. appl. Biol., 33 (in the press).213. TOPPER, B. F. 1942. "Tomato Cultivation on the .College Farm",Trop. Agriculture, Trin., 19, i8o-a.214. TROTTER, A. 1902. "Nuovo contribute alia conoscenza degli Entomocecididella Flora italiana", Riv. Patol. veg., Portici, 9, 359-82.215. 1903. "Nuovi Zoocecidii della Flora italiana", Marcellia, 2,7-23.216. 1904. "Nuovi Zoocecidii della Flora italiana. Seconda serie",Marcellia, 3, 5-13.217. 1906. "Nuovi Zoocecidii della Flora italiana. Quinta serie",Marcellia, 5, 111-23.218. 1907. "Nuovi Zoocecidii della Flora italiana. Settima serie",Marcellia, 6, 102-7.219. TuLLGREN, A. 1917. "Skadedjur i Sverige &ren 1912-1916", Medd.CentAnst. Forsoksv. Jordbr., Stockh., no. 152, 104 pp. (p. 84).220. URICH, F. W. 1915. "Cassava Insects", Bull. Dep. Agric. Trin. Tob.,14,38-40..221. WALTON, C. L. 1927. "Some Observations on the Swede Midge inN. Wales", J. Minist. Agric, 34, 547-51.222. WARBURTON, C. 1926. "Annual Report for, 1926 of the Zoologist",J.R. agric. Soc, 87, 352-6.

References ' 93223. WATTS, F. 1916. "Experiments at the Botanic and ExperimentStations", Rep. Cgnc- Dep. St. Vincent xgrj-igKj, 5-19.224. WiLDMAN, A. B. H)27. "Mocrolabis corrugans on parsnip". Entomologist,60, 259-60.225. WiNNERTZ,J. iSss/'BeitragzureinerMonographiederGallmucken ,Linn. Ent., Bed., 8, 154-322.226. 1854. "Uebef C. juniperina Linn, und C.pisi nov.s]^", Stettin, ent.^tg., ^S.'aaa-T.227. WoRONiECKA J. 1923. ["The Pests of Agriculture observed in Pulawyand its Surroundings in 1923"], Mem. Inst. nat. polon. £con. rur., 4,341-59.228. YuASA, H. & Kui>iAZAWA, T. 1937. ["Distribution and Food Plantsof Asphondylia sp-, a Pest of Soy Bean"], Bot. & ôoL, Tokyo, 5,1561-2.

Index of Generic, Specific apd iôpular Namesof Gall Midges mentionedAertgallmyggen, see pbi Winnertzalpestris Kieffer, (Dasjmeura), 71Amerhapha gracilis Rubsaamen, see under gracilis RuljsaamenArabis Midge, see alpestris Kiefferasparagi Tavares, {Dasyneura), 24, 25Asphondylia ervi Riibsaamen, see under ervi Riibsaan^gulathyri Rubsaamen, see under lathyri Riibsaamen (Asphondylia)sp. on Cassia mimosoides, 59• sp. on Desmodium podocarpum var. japonica, 59-jpi î" JjCgînlktv^ j?^, J57sp. on Glycine ussuriensis, 59sp. on Indogofera pseudotinctoria, 59sp. on Lespedeza bicolor, 59sp. on Soy bean, 59trabuti Marchal, see under trabuti Marchalhamesi Edwards, (Mycophila), 23, 61-2beccariella Del Guercio, {Dasyneura), 22, 25-6Bitter Gourd Vine gall fly, seefalcata FeltBladder Pod Midge, see brassicae Winnertzbrachyptera Kieffer, (JPezomyia), 63brasiliensis Rubsaamen, {Jatrophobia), 22, 30, 31, 32-!}Brassica Flower Midge, see raphanistri KiefferPod Midge, see brassicae Winnertzbrassicae Wiimertz, (Dasyneura), 21, 22, 35, 51-S, 57btyoniae Bouche, (Jaapiella), 21, 23, 80-2Button-top Midge of Willows, see heterobia H. Loewcarophila F. Loew, {Lasioptera), 22, 26Carrot Midge, see pimpinellae F. LoewCecidomyia cucurbitae Felt, see under cucurbitae Felt, {.Oecidomyia)manihot Felt, see under manihol Felt, (Cecidomyia\podagrae Bs, see under podagrae Bssp. on Yam, 82Cecidomyie des fleurs de Cruciferes, 55du chou-fleur, see nasturtii KiefferCecidomyiella crataevae Mani, see under crataevae Mani^ ICecidomyiella)cephalandrae Mani, (Neolasioptera), 22, 33-4cinerea Macquart, (Lestremia), 23, 61, 64Clinodiplosis hydrangeae Barnes, see under hydrangeae B^ês• lathyri Rubsaamen, see under lathyri Riibsaamen (Clinodiplosis)pisicola Barnes, see under/itstco/a Barnessp. in Brassica siliquas, 54• sp. on Ipomoea sp., 77—- sp. on Leek, 23, 57-8sp. on Lettuce, 23, 59-60Colza Flower Midge, see raphanistri Kieffer94

Midgelndex 95Contarinia florum Rubsaamea, see under ^ôrunt Rubsaatnen• gallaica Tavares, see under gallaica Tavareslycopersici Felt, see under lycopersici Feltnasturtii KiefFer, see under nasturtii Kieffer—'— pastinacae Rubsaamen, see under pasiinacae Riibsaamenpisi Winnertz, see under pisi Winnertzpontevedrensis Tavares, see under pontevedrensis Tavaressolani Rubsaamen, see under solani Rubsaamensp. on Brassica, 50sp. on Lactuca virosa, 60tritici Kirby, see under tritici Kirbycorrugans F. Loew, {Macrolabis), 21, 23, 64-7cratojevae Mani, (CecitforrvyteHa), .22, 34, {Neolasioptera), 22, 34cucurbitae. Felt, [Cecidomyia), 23, 77, {Mycodiplosis), 77Dasyneura alpestris Kieffer, see under alpestris Kiefferasparagi Tavares, see under asparagi Tavaresbrassicae Winnertz, see under brassicae Winnertzbeccariella Del Guercio, see under beccariella Del Guercionasturtii Rubsaamen, see under nasturtii Rubsaamenpyri Bouch6, see under pyri BouchS• sp. on Camelina, 53• idriuymOT KiesSfer & Tsystt-sir, see uadsr i&risvcAm KieS^r •& Trottecviciae Kieffer, see under viciae Kiefferdauci Bremi, see pimpinellae F. Loewdizygomyzae Barnes, {Profeltiella), 26Erbsengallmiicke, see pisi Winnertzeni Rubsaamen, (Asphondylia), 23, 58, 59falcata Felt, (Lasioptera), 22, 2Sflorum Rubsaamen, {Contarinia), 22, 24fungicola Felt, {Mycophila), 23, 61, 6?-3gallaica Tavares, [Contarinia), 49-50geisenheyneri Rubsaamen, see nasturtii KiefferGeodiphsis ranunculi Kieffer, see under rammculi Kieffer, (Geodifilosis)Gephyraulus raphanistri Kieffer, see under raphanistri Kieffergracilis Rubsaamen, (Amerhapha), 29Gulerods-galmyggen, see pimpinellae F. Loewheroclei Kaltenbach, see corrugans F. LoewHernesaaski, see pisi Winnertzheterobia H. Loew, (RAabdophaga), 71hieracii Kieffer, [Macrolabis), 67hydrangeae Barnes, [Clinodiplosis), 58inquilina Rubsaamen, [Trotteria), 29ipomoeae Felt, [Schizomyia), 23, 77Jaapiella bryoniae Bouche, see under bryoniae BoucheJatrophobia brasiliensis Riibsaamen, see under brasilierisis Rubsa.amenjavanica Kieffer & Docters Reijnvaan [Prolasioptera), 25Kaalinkukkasaaski, see raphanistri Kieffer,Kamptodiplosis reducta Felt, see imder reducta FeltKiefferia pimpinellae F. Loew, see under pimpinellae F. LoewKohlgallmucke, see brassicae Winnertz *Kohlschotenmucke, see brassicae Winnertz

96 Gall Midges of Economic ImportanceKoolzaadgalmug, see brassicae WinnertzKrauselgallmucke, see nasiurtii KiefferKrusesyge-Galmyggen, see nasturiii Kieffer \lactucae Felt, [Lasioptera), 60 1lamii Riibsaamen, [Macrolabis), 67 1 ILasiopiera carophila F. Loew, see under carophild F. Loewfalcata Felt, see under fakata Felt 1laclucae Felt, see under lactucae FeltLasiopterine Midge on Yam, 82Lasiopteryx manihot Felt, see under manihot Felt, (JLasiopteryx)lathyri Riibsaamen, (Asphondjlia), 26, [Clinodiplosis), 74Leaf-Curling Pear Midge, see pyri BoucheLestodiplosis pisi Barnes, see under pisi BarnesLestremia cinerea Macquart, see under cinerea Macquartlycoperski Felt, (Contarinia), 21, 23, 57, 62, 78-80Macrolabis corrugans F. Loew, see under corruga/is F. Loewhieracii Kieffer, see under hieracii Kiefferlamii Riibsaamen, see under lamii Rubsaamenmanihot Felt, (Cecidomyia),-22, 30, 31, 33, (Lasiopteryx), 22, 30-1manihoti Tavares, (Schizomyia), 23, 30, 31Miastor sp. on Mushroom, 23, 61, 63Mohrengallmiicke, see pimpinellae F. Loewmosellana G^hin, (Sitodiplosis), 71Mycodiplosis cucurbitae Felt, see under cucurbitae Felt, [Mycodiplosis)Mycophila barnesi Edwards, see under bamesi Edwardsfungicola Felt, see under/unîco/a Feltspeyeri Barnes, see under speyeri Barnesnasturtii Kieffer, (Contarinia), 21, 22, 35-51, 53, 55, 6aRiibsaamen, (Dasyneura), 35Meolasioptera cephalandrae Mani, see under cephalandrae Manicrataevae Mani, see under crataevae Mani, (Jêolasioptera)orientalis Felt, (Profeltiella), 26pastinacae Riibsaamen, (Contarinia), 23, 64Pea Midge, see pisi WinnertzPear Leaf Midge, see pyri Bouchepericarpiicala Bremi, see pimpinellae F. Loewperniciosa Riibsaamen, see nasturtii KiefferPezomyia brachyptera Kieffer, see under brachyptera Kieffervanderwulpi Meijere, see under vanderumlpi Meijerepimpinellae F. Loew, (Kiefferia), 21, 22, 26, 27-30pisi Barnes, (Lestodiplosis), 74, 76Winnertz, (Contarinia), 21, 23, 68-76pisicola Barnes, (Clinodiplosis), 7a, 74, 76Meijere, see pisi WinnertzPod Midge, see brassicae Winnertzpodagras Bs, (Cecidomyia), 60pontevedrensis Tavares, (Contarinia), 49, 50Profeltiella dizygomyzae Barnes, see under dizygomyzae Barnesorientalis Felt, see under orientalis Feltranunculi Kieffer, see under ranunculi Kieffer, (Profeltiella)• soya Monzen, see under soya Monzensp. on Broad Bean, 22, 26sp. on Rhubarb, a3,*77 ,vespicoloris Barnesj see under vespicoloris Barnes

Midgelndex 97Prolasiopiera javanica Kieffer & Docters Reijnvaan, see under javanica Kicffer &Docters Reijnvaanpyri Bouche, [Dasyneura), 71ranunculi Kieffer, (Geodiplosis), 26, {Prqfeltiella), 26raphanistri Kieffer, (Gephyraulus), 22, 53, 55-7reducta Felt, {Kamptodiplosis), 26Rhabdophaga heterobia H. Loew, see under heterobia H. LoewSchizomyia ipomoeae Felt, see under ipomoeae Feltmanihoti Tavares, see under manihoti TavaresSitodiplosis mosellana Gehin, see under mosellana Gehinsolani Riibsaamen, {Contarinia), 78soya Monzen, {Profeltiella), 26speyeri Barnes, {Mycophila), 23, 61, 63Swede Midge, see nasturtii Kiefferthysselini H. Loew, see pimpinellae F. LoewTomato Flower Midge, see lycopersici Felttorquens Meijere, see nasturtii Kieffertrahuti Marchal, (Asphondylia), 23, 76tritici Kirby, (Contarinia), 71Trotteria inguilina Riibsaamen, see under iriguilina Riibsaamenumbelliferarum Kieffer, see under umbelliferarum Kieffertudensis Tavares, (Contarinia), 49turionum Kieffer & Trotter, {Dasyneura), 24, 25Turnip and Cabbage Seed Midge, see brassicae Winnertzumbellatarum F. Loew, see pimpinellae F. Loewumbelliferarum Kieffer, [Trotteria), 28, 29Unidentified Asparagus Midges, 22, 24, 25Bean Midge, 26Bitter Gourd Midge, 22, 25• Coccinia cordifolia Midge, 34Egg-plant Midges, 23, 57Lentil Midge, 23, 59Rhubarb Midge, 77Yam Midges, 23, 82vanderwulpi Meijere {Pezomyia), 23, 61, 63Vegetable Marrow Midge, see bryoniae Bouchevespicoloris Barnes, (Prqfeltiella), 26viciae Kieffer, (Dasyneura), 23Wheat Blossom Midges, see mosellana Gehin and tritici KirbyWhite Bryony Midge, see bryoniae Bouche

Plant IndexAEPIM, 22Agaricus arvensis Qu^l, 60-4campestris Quel, 60-4Allium porrum L., 23, 57-8Angelica sylvestris L., 28, wild, 28Anthriscus silveslris Hoffm., 28ASPARAGUS, 22, 24-SAsparagus acutifolius L., 24albus, L., 25aphyllus L., 24horridus L., 25officinalis L., 23, 24AUBEROINE, 22, 57Balmoral, variety of swede, 45Bangholm, variety of swede, 45BEAN, BROAD, 22, 26, 72Best of All, variety of swede, 45BITTER GOURD, 22, J5-6Brassica adpressa Bois, 56• amplexicaulis Janka, 45—— campestris L., 45, 53, 56cheiranthus Vill., 53fruliculosa Cyrill spp. radicata Dsf.,50napus L., 22, 38, 45,49, 50, 53,56oleracea L., 22, 45, 53, 56rutabaga, 45, 53, 56spp., 40, 45, 53, 56BROAD BEAN, 22, 26, 72Broccoli, 35, 45Bryonia alba L., 80, 81(fioiVa Jacq., 80, 81Bryony, White, 80, 81Buffalo, variety of s\vede, 45Bupleurum falcatum L,, 28longifolium L., 58rammculoides L., 28Buplever, Sickle-leaved, 28Burnet Saxifrage, 28CABBAGES, 22, 34-5?Cabbage, Cow, 35, 36, 45, Jersey, 53, Savoy, 35, 36, 45Camelina sativa Cr., 53Caraway, 26, 28CARROT, 22, 26-9, wild, 27Carum carvi L., 28CASSAVA, BrrxER, 22, 30, SWEET, 22, 30Cassia acutifolia Delile, 59mimosoides, 59CAULIFLOV^TER, 22, 36, 37, 4a, 44, 45,47. 49Chaerophyllum aromattcum L., 66hirsutum Vill., 66Charlock, 45, 53Chervil, 28, 66Clover, Red, 58Coccinia cordifolia Cogn., 34iNDicA (Naud.) Wight & Arn., 22,33-4Cochlearia armoracia L., 45Colza, 22, 51, 55, 56Cosmos, 58Cow Cabbage, 35, 36, 45parsnip, 28, 66CBATAEVA BBLIOIOSA ForSt., 22, 34CRUCIFERS, CULTIVATED, 22, 34-57, wild, 53Cucurbita maxima, 77pepo L. var., 23, 77, 80Daucus carota L., 22, 26-9var. proslratus Rouy &Camus, 28Dead-Nettie, Red, 67, White, 67Desmodium gyrans DC, 59podocarpum var. japonica, 59Dioscorea minutiflora Engl., 82spp., 23, 82Dreadnought, variety of swede, 45Eclipse, variety of swede, 45EGG-PLANT, 23, 57, 79Erucastrum incanum, Koch, 49pollichii Schimp. & Spann., 56Ervum lens L., 58Falcaria rivini'Host, a8Fennel, 28Foeniculum officinale AH., 2898

Foeniculum piperitum Sweet, 28Gladiolus, 58Gladstone, variety of pea, 72Glycine max (L.) Merr., 58ussuriensis, 59GOURD, BITTER, 22, 25-6Hawkweed, Savoy, 67Hedge Woundwort, 67Hedge-parsley, Upright, 28Heracleum sphondylium L., 28, 65, 66, 67Hieracium boreale Fries, 67Hog's Fennels, 66Hogweed, 28, 66Hollow Crown, variety of parsnip, 65Hollyhock, 58Horse-radish, 45Hydrangea, 58Indigqfera pseudotinctoria, 59Ipomoea batatas Lam., 23, 77^ sp., 77Jersey Cabbage, 53Kale, 35, 36, 45Lactuca sativa L., 23, 59-60virosa L., 60Lamium album L., 67flexuosum Ten., 67• maculatum L. var rugosum Briquet,67pwrpureum L., 67spp., 67Laserpitium pruthenicum L., 28Lathyrus odoratus L., 72, 74Leader, variety of pea, 72LEEK, 23, 57-8Lens esculenta Moench, 23, 58-9LENTIL, 23, 58-9Lespedeza bicolor, 59striata Hook. & Am., 59LETTUCE, 23, 59-60——, Acrid, 60, wild, 60Lobelia, 58Lord Derby, variety of swede, 45Magnum Bonum, variety of swede, 45MANDIOO, BITTER, 22, 30-3SWEET, 22, 30-3Manihot aipi Pohl, 22, 30-3dichotoma Ule, 32palmata Mtill.-Arg., 23, 30, 32utilissima Pohl, 22, 30-3Maranta arundinacea, 30Meadow Saxifrage, Annual, 28Melothria perpusilla Cogn., 25Model, variety of swede, 45Mamordica charantia, 25pterocarpa Hochst., 25Plant Index 99Momordica pteromorfa, 25spp., 22, 25-6Monarch, variety of swede, 45MUSHROOM, 23, 60-4Mustard, black, 53, white, 53, wild, 45Nasturtium amphibium, 45officinale R- Brown, 45palustre DC, 38, 40, 45pyrenaicum R. Brown, 45silvestre R. Brown, 45spp., 45Nightshade, Woody, 80Parsley, 28PARSNIP, 23, 28, 64-7, variety of. Hollow Crown, 65Pastinaca sativa L., 23, 64-7PEA, 23, 68-76, FIELD, 72, GREEN, 72, Sweet, 72, 74, variety of, Gladstone, 72, , Leader, 72, , Senator, 72PEPPER, 79Petroselinum sativum Hoffm., 28Peucedanum cervaria Lapeyr., 66oreoselinum Moench, 28, 29, 66palustre Moench, 28Physospermum aquilegifolium Koch, 28Pimpinella magna L., 28saxifraga L., 28tragium Vill., 28Pisvm arvense L., 23, 72sativum L., 23, 72POTATO, 23, 76Psaliota arvensis Quel, 23, 60campestris Quel, 23, iSoPUMPKIN, 77• , giant, 77RADISH, 22, 23> 35> 53. 56, wild, 45, 56Ranunculus acer, 26RAPE, 22, 23, 35, 36, 45. 51. 53. 56Raphnnus caudatus L., 45, 56raphanistrum L., 45, 56—— sativus L., 43, 53silvestre Asch., 49, 56silvestris Lam., 50Red Clover, 58Red Dead-Nettie, 67Rheum rhaponticum L., 23, 77RHUBARB, 23, 77Savoy cabbage, 35, 36, 45Hawkweed, 67Senator, variety of pea, 72SEPIM, 23'

011>S51100 Gall Midges of Economic ImportanceSesili annimm L., 28(Libanotis) montana, 29 •Silaus pratensis Besser, 28Sinapis alba L., 53arvensis L., 45, 53—— nigra L., 53Siphonodon celastrineus, 26Solanum dulcamara L., 80lycopersicum L., 23, 78-80• melongma L., 23, 57tuberosum L., 23, 76Soy bean, 58SQUASH, 23, 77Stachys sylvatica, 67Superlative, variety of swede, 45SWEDE, 22, 23, 45, 53, 56, variety of, Balmoral, 45—, Bangholm, 45—, Best of Ail, 45—, Buffalo, 45—, Dreadnought, 45—, Eclipse, 45—, Lord Derby, 45—, Magnum Bonum, 45—, Model, 45Swede, variety of, Monarch, 45, , Superlative, 45Lj ^ Wilhelmsberger, 46Sweet Pea, 72, 74SWEET POTATO, '23, 77TOMATO, 23, 78-80Torilis anthriscus Gmelin, 28TURNIP, 22, 23, 45, 53, 56VEGETABLE MARROW, 23, 80-2Vicia [Ervum) hirsuta Koch, 59• tetrasperma Moench, 59faba L., 22, 26Watercress, Marsh, 38, 45, Creeping, 45White Bryony, 8x, 82Dead-Nettie, 67Wilhelrnsberger, variety of swede, 46willow, rotten, trunk, 63Woody Nightshade, 80Woundwort, Hedge, 67YAM, 23, 82

General IndexAepim, midge on, 22aestivation, 71ammoniiun oleate, 48valerianate, 48Anthocoris nemorum, as enemy of Mdcrolabiscorrugans, 67Aprostocetus fidius, as parasite of Jatrophobiabrasiliensis, 33sp., as parasite of Jatrophobiabrasiliensis, 33apterous forms, 6$arrowroot, 30artificial manures, 47, 67Asparagus, midges on, 22Aubergine, midges on, 22.bacterial rot, associated vvfith Contarinianasturtii, 36Barnes slide collection of gall midges,14-15Beans, midges on, 22binodal curve of emergence, 71, 73biological control of Contarinia pisi, 76Bitter Cassava, midges on, 22Gourd, midges on, 22"bladder pod", 51blindness in savoy cabbage, 39in cauliflower, 36Brazilian arrowroot, 30Broad Bean, midges on, 22Cabbages, midges on, 22"cabbage-top" of swedes, 35, 38calcium cyanamide, 47carbol-turps preservative mixture, 13Carrot, midge on, 22cassareep, 30Cassava, midges on, 22Cauliflower, midge on, 22Ceuthorrhynchus assimilis, 52-3Chalcid parasite of Dasyneura beccariella,62Jaapiella bryoniae, 82Macrolabis corrugans, 67Chemical methods of control :ammonivrai oleate,,48valerianate, 48calcium cyanamide, 47chloronaphthalene, 48Chemical methods of control:clay as a drying agent, 62creosote, 48derris, 48, 49ethyl hydrosulphide, 48ground nut oil, 48kainit, 47, 75lime, 47as a drying agent, 62methylated spirits, 48naphthalene, 48, 74, 79nicQtine, 49, 74, 79, crude, 48, sulphate, 49, 74nitrate of soda, 47oleic acid, 49paradichlorobenzene, 47, 79para-phenylenediamine, 48pyrethrum, 48, 62, 80pyridine, 48, 49sodium sulphoricinate, 49soft soap, 48, 49, 74sylvinite, 47Vaporite, 79volatile alkali, 49chloronaphthalene, 48clay as a drying agent, 62clean cultivation, 46Coccinia indica, midge on, 22coco-nut fibre for soil-inhabitinglarvae, 10Colza, midges on, 22Contarinia lycopersici Felt:closely allied species, 80control measures, 79damage, 78description, 78diagnostic character, 78distribution, 78food plants, 79life history, 78material, 80parasites, 79references, most important, 80Contarinia nasturtii KiefTer:closely allied species, 49control, cultural methods, 46to kill larvae in the soil, 47repellents, 47101

102 Gall Midges of Economic importanceContarinia nasturtii Kieffer (contd.):control, sprays, 48damage, 35description, 38diagnostic character, 35distribution, 41fluctuations in numbers, 36food plants, 45life history: „in Denmark, 41, 44• England, 41-4France, 41-4Germany, 41-3, 45Holland, 41-4material, 50natural enemies, 46references, most important, 50resistant varieties, 45synonymy, 38lontarinia pisi Winnertz:biological control, 76control, 74damage, 68'description, 69diagnostic character, 68distribution, 69fluctuations in numbers, 72food plants, 72inquilines, 74life history, in England, 70-1, in Switzerland, 70-1material, 76natural enemies, 73references, most important, 76resistant varieties, 72synonymy, 69:ontrol, see biological controlsee chemical methods ofcultural methods ofhot waterIrataeva religiosa, midges on, 22;reosote, 48Titeria for systematists, 10:rop rotation, 46, 54, 75:rops, list of, 22-3Cruciferous crops, midges on, 22'crumpled-leaf", 35, 38, 40;ultural methods of control:artificial manures, 47, 67clean cultivation, 46crop rotation, 46, 54, 75deep digging, 75•—— ploughing, 54early varieties, selection of, 75handpicking, 54, 62, 75, 82late sowing, 47natural barriers, 75nip out terminal buds, 82ploughing, 72, 75trap crops, 46, 76'curly-leaf", 35damp soil necessary for pupation, 79Dasyneura brassicae Winnertz:I closely allied species, 55control, 54damage, 51, description, 51'.diagnostic character, 51distribution, 51'food plants, 53inquiline, 54life history, 51-2material, 55natural enemies, 54references, most important, 55resistant varieties, 54deep digging, 75ploughing, 54delayed emergence, 9, 43, 71depth of pupation in soil, 43, 52, 70,79derris, 48, 49"draaihartigheid", 39"drehherzigkrankheit", 39drought, effect of, on larvae, 42dry rot, 36—— season, incidence of Contarinialycopersici in, 79duration of adult midge stage, 12, 70egg stage, 34, 41, 66, 70, 78larval stage, 34, 42, 66, 70, 79, 81pupal stage, 34, 43, 70, 79, 81"early" damage of tomato fruit, 78sowing, 75varieties, selection of, 75egg laying oi Dasyneura brassicae, 52Egg-plant, midges on, 23emergence, binodal curve of, 71, delayed, 9, 43, 71Empid fly, as enemy of Contarinianasturtii midges, 46ethyl hydrosulphide, 48fibre for soil-inhabiting larvae, 10fluctuations in numbers, 36, 68fungal infection, associated with Contarinianasturtii, 36fungus attacking Dasyneura brassicae, 54gall midges, list of, 22-3, monographs on, 7galls, comprehensive works on, 7generation, definition of a, 70Gephyraulus raphanistri Kieffer:closely aUied species, 57control, 57damage, 55description, 56diagnostic character, 55distribution, 56food plants, 56^life history, 56material, 57

denefal Index 103Gephyraulus raphanistri KiefFer (contd.):natural enemies, 56reference, most important, 57ground nut oil, 48handpicking, 54, 62, 75, 82heliotropism, 61hibernation, 11, 28, 43, 53, 56, 70,81host plant range of gall midges, 9,67,79hot water control, 63humidity, effect of, on prevalence oflarvae, 38, 42, 53, 79Inostemma boscii, as parasite of Contariniapisi, 73-4, as parasite oiDasyneura brassicae,54inquilines:Amerhapha gracilis, 29Clinodiplosis pisicola, 74sp. with Dasjneura brassicae,54 .sp. possibly with Contartmasp. on lettuce, 59-60Trotteria inquilina, 29umbelliferarum, 28-9Jaapiella hryoniae Bouche:control, 82damage, 80description, 80diagnostic character, 80distribution, 81food plants, 81life history, 81material, 82natural enemies, 82references, most important, 82kainit, 47, 75keys for the separation of genera, 9Kiefferia pimpinellae F. Loew:control, 29damage, 27description, 27diagnostic character, 27distribution, 27food plants, 28inquilines, 28-9life history, 28material, 29natural enemies, 28references, most important, 30synonymy, 27"krauselkrankheit", 39labelling, 14larval development, retardation of, 9,late sowing, in preventmg attack, 47Leek, midge on, 23Lentil, midges on, 23Leptacis tipulae, as parasite of Contariniapisi, 73Lestodiplosis pisi, as predator of Contariniapisi, 74Lettuce, midge on, 23lime, 47âs a drying agent, 62Macrolabis corrugans F. Loew:control, 67damage, 65description, 65diagnostic character, 65distribution, 66food plants, 66life history, 66material, 67natural enemies, 67references, most important, 67mandioca flour, 30Mandioc, Bitter, midges on, 22, Sweet, midge on, 22manures, artificial, 67"many-necked" condition of swedes,35> 38method of labelling slides, 14larval feeding, 42, 52recognizing Barnes cotypes, 15homotypes, 15paratypes, 15 . •types, 14storing slide collection, 14methods for breeding gall midges, 10-12mounting empty pupal cases, 13larvae, 13larval skins, 13midges, 13preserving larvae, 12midges, 12pupae, 12methylated spirits, 48Mushroom, midges on, 23muslin cages, 10covered iron rings, 10naphthalene, 48, 74, 79natural barriers, 75nematode killing midge larvae, 63"nettlehead" of peas, 68nicotine, 49, 74, 79, crude, 48, sulphate, 49, 74nip out terminal buds, 82nitrate of soda, 47number of generations, 28, 34, 43, 53,56, 66, 70, 81numbers of larvae per acre, 69gall, 24, 25, 52,63, 66, 68, 81oleic acid, 49paedogenesis, 61-3

104 Gall Midges of Economic Importancebecparadichlorobenzene,47, 79para-phenylendiamine, 48parasites:Aprostocetus fidhis, 32-3—— sp., 33Chalcid parasite of Dasyneuracariella, 26Jaapiella bryoniae, 82Macrolabis cormgans, 67I'Inostemma boscii, 54, 73-4Leptacis tipulae, 73nematode killing midge larvae, 63Pirene graminea, 73-4, 76Platygaster cruciferarum, 56sp. (? nigra), 54Proctotrupid parasites of Contannianasturtii, 46Prosactogaster floricola, 56Pseudotorymm brassicae, 54Sactogaster pisi, 73sp., 79Synopeas daucicola, 28raphanistri, 56Tetrosiichus brevicornis, 54-fasciatus, 31-3Parsnip, midges on, 23Pea, midges on, 23 .peat for soil inhabiting larvae, 10Phoma Lingam, 36Phototropism, 62Pirene graminea, as parasite of Contariniapisi, 73-4, 76Platygaster cruciferarum, as parasite ofGephyraulus raphanistri, 56sp. (? nigra), as parasite of Dasyneurabrassicae, 54ploughing, 72, 75Potato, midge on, 23,- S^yeet, midjre on, 33predators:Anthocoris nemorum, 67Empid fly, 46Lestodiplosis pisi, 74spiders, 46preservative mixture, 13Proctotrupids, as parasites of Contarinianasturtii, 46Prosactogaster Jloricola, as parasite ofGephyraulus raphanistri, 56Pj. destructans, 36PseudotOrymus brassicae, as parasite ofDasyneura brassicae, 54pyrethrxun, 48, 62, 80pyridine, 48, 49Radish, midges on, 23Rape, midges on, 23rate of unfolding of leaves, 64-5retardation of larval development, 9,43.71Rhubarb, midge on, 23Sactogaster pisi, as parasite of Contariniapisi, 73sp. as parasite of Contarinia lycopersici,79senna, 59Sepim, midge on, 23sJide cbJJectJDn, method of storing theBarnes, 14sodium sulphoricinate, 49soft soap, 48, 49, 74sparrows, precautions against, 11spiders, as enemy of Contarinia nasturtiimidges, 46Squash, midge on, 23stage of grovvrth of plant for successfulinfestation, 36, 42, 64, 73submersion, resistance of larvae to 4.2,62sunlight, effect of direct sunlight oneggs and young larvae, 65Swedes, midges on, 23Sweet Cassava, midge on, szPotato, midge on, 23sylvinite, 47Synopeas daucicola, as parasite of Riefferiapimpinellae, 28raphanistri, as parasite of Gephyraulusraphanistri, 56tapioca, 30Tetrastichus brevicornis, as parasite ofDasyneura brassicae, 54fasciatus, as parasite of CeciJomyiamanihot, 31, Jatrophobia brasiUensis,32-3Tomato, midge on, 2gtrap crops, 46, 7,6Turnip, midges on, 23Vaporite, 79Vegetable Marrow, midge on, 23volatile alkali, 49wasps, precautions against, 12water, resistance of larvae to, 42, 62weather, effect of, on incidence ofContarinia nasturtii, 37-8weeds as alternate food plants, 28, 46,55>66weevils, presence of, necessary forsuccessful oviposition of Dasyneurabrassicae, 52-3wet season, incidence of Contarinialycopersici in, 79Yam, midges on, 23Printed in Great Britain by Butler St Tanner Ltd., Frome and London

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GALL MIDGES ECONOMIC IMPORTANCE - KrishiKosh

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