J. ESTU\I

TilE MARSH CRA~E FLY, TJPULA PALUDOSA Mg., A NEW PEST IN BRITISH COLLMBIA (DIPTERA: TIPULIDAE)

A. T. S. WILKINSON AND H . R. MACCARTHY I ABSTRACT Tipula paludosa. Meigen was firmly established in the Vancouver area by 1965. startmg In the eastern outskirts of thc city. It was taken at Blame, Wash.,. 111 1966. By 1967 thc pcst had spread to thc Chilliwack area P opulatlOns of 110 ftz were mcasurcd in 1966 and lucal observatiuns \\'c rr recorded of damage, oviposition, fecding, growth, and cmel'gen~e. A revIew IS mcluded of Europcan literaturc on populations life history weather relatlOns, and blOlogical, chemical and cultural ~ontrols with some spcculatlOn on the future. '

INTRODUCTION damage in 1959 to cabbage trans­ In the past three years the leath­ plants and turnip seedlings. The or­ erj acket or larva of the European igin of the pre sen t outbreak is a marsh c ra n e fly, Tipula paludosa matter for speculation; a good guess Mg., has become a serious problem in would be the balled roots of orna­ lawns and pastures in the lower Fra­ mentals imported from Europe. ser Valley. Identification was by J . R. LOCAL OBSERVATIONS Vockeroth, Entomology Research In­ Mos t of the damage in British Co­ stitute, Ottawa. This is the most com­ mon and damaging crane fly in lumbia has been to lawns and pas ­ tures but flowers, strawberries and northwestern Europe. The leather­ H'getable crops in backyard gardens jackets were first found in 1965 caus­ ing severe damage to lawns in the have also been attacked. These infes­ eastern outskirts of Vancouver. In ta tions have been easily con trolled with DDT or aldrin, but the rapid 1966 they were considerably more spread of this pest to pasture has widespread and in pastures on several presented a much more serious prob­ small farms in this area there was lem. The lower Fraser Valley is pri­ virtually no growth until the middle marily a dairying region, and there of May. In the fall adult T. paludosa is great danger of insecticide residues were trapped at Blaine, Washington, occurring in meat and milk fats if 25 miles sou the as t of Vancouver (U.S.D.A. Cooperative Economic In­ these and similar persistent insecti­ cides are applied to forage or fodder. sect Rpt. 16: 946, 949, 956. 1966). In 1967 heavy infestations of leather­ In 1966 preliminary stUdies were jackets and the resulting damage carried out in a heavy infestation on occurred on large dairy farms near a lO-acre farm. In April and May the Pitt Meadows, 20 miles east of Van­ larval population was measured in 84 couver, and in lawns at Yarrow 30 samples, each of 1/ 4 sq ft, on about 4 miles farther east. . , acres. Gasoline sprayed on the turf brought nearly all the larvae to the The first North American record surface where they were rea d i 1 Y of this pest was in 1955 on Cape Bre­ counted. The top I-inch of sad was ton Island (Fox, 1957) where lawns examined for the remainder that did and flowers were attacked. The in­ not emerge. The population per sq festation there was thought to have ft averaged 109.6 (range 24-232), or originated in soil used for ships bal­ close to 5 million per acre. last and dumped ashore. In New­ foundland M 0 r r is (1960) reported Rototilling and disking reduced the population by about two-thirds , Contributio n No. 132 Researc h Station, Re· bu t the reduction was not enough to search Branch. Canada Department of Agricul· ture, 6660 N.W. Marine Drive, Vancouver B. B.C. allow a new seeding to survive. When ::n .T. r£"T "~ I()I .. SO('. BlOT. COI .D ITlJA. 64 (1967 ). AUG . 1, 1967 oat." anci gra.<; s were planted on April ea.se in this infestation. 26 nothing grew. not even volunteer grass or weeds. The larvae matured POPULATIONS It and .~ topped feed ing about th e middle is clear th at n umbers of th e of May. Oats a nd grass planted 811 pest wer e in a runaway ph ase during May 20 prOd l! Ccd a good crop. J9 66. Maercks (1939) in northern On war m, cl oudy days in March Germany, Go ns icl ered Lhat 5 per ;;0 and .-1.pril, the larvae were observed I t would cause serious m jury in ar­ on the surface feeding on the crowns a bie land, ancl 10 per sq It serious a nci blades at' grass. On bright days injury in grass land. Coh en & st e l~ r thev remained in thc sod but fed on , 1946) cons idered 20 per sq It to be the .': llrface at !light. The larvae were J, ;1eavy infestation. 'The populatic"l e,enerally fo und in th e LOp I - inch of :evel is likely to decline as exis ti '.' sod . out a lter about ~ he middle of and imported biological controls a'­ May they moved downward and m a ny .c;en themselves. wer e found a.'i deep as 3 inches below LIFE HISTORY Ghe surface. Several expenenced investlga LO r.s "": mergence a nd a dult populations have studied T . {Ja l ucloset, a n d th eIr were studied by placing six l -sq- ft a CCOUl1 ts a r e well :n agr eem ent WIth cages in a pasture a nd taking counts one another . The species a ppears cG (wice weekl y. The ad u 1 t s started be mostly u 11 i v 0 I tin e but :,;ome em ergmg during the fi r.st week of a uthors s tate wi thout offering evi­ August. ;Jeaked a bout September 1, den ce that t h ere may be a par tial or a nci th e last on e was collected on eomplete 2nd generation . Se)J tember 30. The average number Egg. The eggs are black a nd shiny, of adults pel' sq ft \/;,as 98 (range 1.1 x 0.4 mm, J, r e laid at nigh t in 75-112) consis ting of 55 males, 43 fe ­ August and September, and have very males. high moistur e require m en ts at first. The egg capacity was observed by They will collapse within 2 to 4 m in­ Coulson's (1962) m ethod of removing ll tes in less th an 100% relative h umi­ th e head fr om newly- em erged and dity (Laugh lin, 1958) . Th e min imum m ated females from th e field and morta lity occurs in upla nd soil hold­ floating them on water wh ich inducE'S ing t wi ce its d ry weigh t of water th em to oviposit. The eggs remaining (Maercks , 1939). They are laid on or in th eir a bdomens wer e a l.so counted . very close to th e surface, 68% within Th e a verage n umber of eggs based on 1 em of it according to Co u Iso n 10 fem a les was 281 (r a n ge 24 3-338) . (1 962). Ren n ie (1916) considered th at Th is is lower th a n th e average of 360 not a ll th e eggs wer e m ature on th e reported by Coulson (196 2) in th e emergence of th e female, and th at n orth of England but close to Barnes' som e were retained to produce a 2nd (1937) fi gure of 272 in th e south . batch. Most later workers think th a t No paras ites h ave emer ged from rem aining unla id eggs a r e simply a many hundreds of la r vae r ea red in residue. The eggs develop with ou t Gh e laboratory, n or have larvae been diapa use (Sellke, 1936). observed killed by virus. The only L arva . The larva e h a tch in 11 to predators seen we r e sp iders and pos­ 15 days (Rennie, 1917 ; Ba rnes, 1937) . sibly th e n umerous European star­ They feed from the first day, s ta rting li n gs. Sturllu s v u lgaris, which a ppear a t a pproximately 2. 7 mm long, with to feed on larvae a nd a d ul ts. Aver age 13 segments, growing to 4 or 5 mm survival from th e late larval stages to in 12 to 13 days (Rennie, 1917 ). Mor ­ a d ults was high wh en 98 adults we re tali ty is high for th e first 20 to :30 obtained from an average of 11 0 l3. r­ days (La ughlin, 1958) . vae per sq ft . This s uggests th at th ere The first two inst a rs of three are are at presen t low levels of ca nni­ passed in a bout 14 days in centra l balism, preda tion, parasitism a nd dis - Engla nd (Ba rnes, 1937). In six weeks J. ENTOMOI.. SOC. HRlT. COI.l T ~mIA. 64 (1967). ArG. 1. 1967 31

they a re about 6 mm long. The winter 196 ~J and have finish ed laying within is spent in the 3rd instar (Fig. 1), 32 hrs of emer ging (Coulson, 1962). which is without diapause a nd lasts They fly very little before laying, but roughly 25 weeks (Sellke, 1937; Coul­ m a y leave 5 or 6 eggs in one spot, son, 19 62 ) . Growth is r apid in spring th en move a short distance (Rennie, when m ost of th e damage occurs 1917 ) . Males live about 7 days, fe­ (Rennie, 1917). Minimum larval mor­ ma les from 4 to 5 days (Barnes, 19 37) . tality occurs in upland soil holding In a 4-year study at Roth amsted, three tim es its dry weight of water 57 % of 3,400 a dult crane-flies were (Maer cks. 1939 ) . Yo ung larvae appear of th is species (Robertson, 1939). to prefer gr een leaves to roots and grow lDus t rapidl/ with least los.., JC WEATHER RELATIONS white clover . They a lso rear easily on Maercks (1941) concludes th at lettuce, wh eat and rye, but m ay not this pest is favored by mild winters, complete d eve l op men t on oats " 001 snmmen , and rainfall averaging (Maercks, 19 39). Normally, but not at least 24 inch es per year. Mean invariably, they s urface to feed, dur ­ monthly values for temperature, pre­ ing darkness (Sellke, 1937). cipitation a n d numbers of days with ra infall are s how n by months in Pupa. The la rvae stop feeding in Ta ble 1, based on 3D -year averages mid-May for two or three weeks be­ at th e Vancouver Internationa l Air­ fore pupating (Fig. 2). W h en the port. Values for agricultural a,'eas of a dult is ready to em erge th e pupa the lower Fraser Va lley differ only works its way to the surface where s lightly. It thus appears that th e the empty pupal case is left by th e maritime climate of the wet, coastal emerging a d u I t, ch aracteristically belt of British Columbia is practically protruding 2. 5 cm from th e ground. idea l for this pest. Th ese a r e easily sexed, and a ratio of 1.72 m a les to 1 female was establish­ The prespn t au tbreak h as been <: d by Coulson (1962) . Cannibalism favored by l' ecent weather patt.erns. may be a r educing factor in th e early In the 5 yea rs , 1962-1966, mean stages of pupation. monthly temperatures in winter and summ er were a bove normal as fol ­ Adult. The adults emerge soon lows: after sunset in August a nd septem­ Nov. Dec. J an. Feb. May ber, m ate immediately (Fig. 3), lay 4/ 5 2/ 5 3/ 5 5/ 5 0/ 5 75 % of th eir eggs before daylight June July Aug. (Sellke, 19 37; Ba rnes, 1937; Coulson, 0/ 5 0/ 5 0/ 5 The conclusion is that the pest h as TABLE I-Mean monthly temperatures, h ad five years of idea l conditions in precipitation, and days with rain, based on 30 years of r ecords, Vancouver Inter national which to become well establish ed . Airport. Da m age may be expected follow­ Mean mo. Mean mo . Mean days ing a wet September, especially if th e Month temp.," F precipitation, with rain following winter is mild (Maercks, inches 1941) . A cold spring contributes to Jan. 37.2 5.52 19 Feb. 39.4 4.74 16 damage, beca use the danger p eriod in March 43.2 3.76 16 annual crops is from th e time of sow­ April 48 .3 2.30 13 ing to the growth of adventitious May 55.0 1.92 10 roots (R e n n i e, 1917 ). Rober tson June 60.4 1.84 9 July 63.8 1.04 6 (1939) noticed that twice as m any Aug. 63.6 1.37 8 a dults were taken at ligh t traps on Sept. 57.8 2.13 9 moonless nights as on moonlit ones, Oct. 50.3 462 15 and three times as many on cloudy Nov. 43.1 5.44 18 Dec. 39.6 6.44 20 as on clear nights. Most of th e trap­ Total 41.12 159 ped adults were males . .J , 1 ~"' III ,\ III i " S, ,, , UH I I' , ('11 1. 1' \ 11: 1,\ , G4 (1967 ) , A UG , 1, 1967

,/ J . E NTOMOL. Soc. BRIT. COLUMBIA, 64 (1967) . AUG. 1, 1967 33

BIOLOGICAL CONTROLS h azard, and how to live with the pest T. paZudosa is not effectively con­ at th e same time keeping down costs. trolled naturally in northwest Eur­ Fortunately the larvae are easily ope. The most effective insect para­ killed and will accept baits readily. site appears to be Siphona genicuZata They have thin in tegument, perme­ De Geer, a small Tachinid that lays a ble enough for gaseous exchange, up to 9 eggs on the stigmatic crown lacking an epicuticular layer (Ghil­ of the leatherjacket. The larvae en­ arov & Semenova, 1957). Moreover, ter the main tracheal trunks and 1st and 2nd instar larvae remain close bore into the h emocoele but retain a to the s urface, and have been killed respiratory connection with a chiti­ even by mineral fertilizer (Sellke, nous sh eath-like structure. There are 1937). two generations per year and the parasite overwinters in the host, but CULTURAL CONTROLS the level of paraSitism is n ever high. Leatherjackets may be reduced by One record shows 34% to have been cultivation, since they do not go deep affected but the average is much low­ into th e soil, but when the numbers er, from 6 to 17% (Rennie & Suther­ are very large the reduction m ay not land, 1920) . The Vancouver Station is be effective. However , T. paZudosa is a ttempting to establish S. geniculata adaptable enough to survive and re­ supplied by the Institute for Biologi­ produce without the pre sen c e of cal Control, Belleville, Onto growing plants, by eating decaying Two virus dis e as e s of leather­ rootlets a fter the manner of wire­ jackets have been recorded and plans worms (R ennie, 1917). M ae r c ks to use these are under way at Van­ (1941) advocates good drainage of couver. However , they do not a ppear land and short grass during egg-lay­ to be highly contagious, although fa­ ing in August and September. The tal. Empusa ( = Entomophthora) has deleterious effect on the eggs and been recorded as infesting popula­ young larvae of dry weather, may tions in Germany (Muller - Kogler , sometimes be offset by the practice 1957), and a fungal infection of the of irrigating pastures with .sprinklers. tracheae is known (Coulson, 1962). FUTURE PROSPECTS Two species of saprozoic and parasitic nematodes have been recorded in Review of th e extensive European Denmark (Bovien , 1937). Cannibal­ literature indicates that T . paludosa ism is a mortality factor in la bora­ is likely to become a constant and tory rearing, but its effect in the field possibly a major pes t in areas of high is difficult to assess and probably rainfall. There a re dozens of records small. George (1966) concludes that of damage by this speCies in north­ there is little evidence for effective western Europe in research papers diseases. and annual reports from Denmark, Predation on larvae, especially by U.K., Germany, Sweden and Holland. European starlings and na tive moles, It probably will establish itself in northwestern Washington, but its Scapanus spp., should be studied. Predation on adults is probably not south ern a nd eastern spread may be effective, since any adult taken is restricted by cold winters and by its likely already to have expended most high moisture requiremen ts. Popula­ t io n crashes in northern England in of its quota of eggs. 1955 and 1959 were shown experi­ CHEMICAL CONTROLS men tally by Milne et al. (1965) to re­ The problem is twofold . .ow to sult from very dry conditions at criti­ treat land without creating a residue cal periods. Fig. I-Mature 3rd-instar larvae of T. paludosa. Fig. 2- Pupa of T. paludosa. Fig. 3-Marsh crane flies, Tipula paludosa in copula. 34 .J. [<;"T""I>I . SI>,. B lnT. (" " . I · ~ I "I..\. f;~ (1f167l. Ann. I , J 967

Th ey predicted th at in creases may Nevertheless th ere is likely always to be expected if rainfall in August a nd be a res idue in low-lying land and September is normal or greater than in ditch-banks. norma l, but wh en rainfall at this It appears unlikely th at r esistance time drops below 50 % of normal, to ch emical pesticides will develop sh a rp declines will certainly occur. within 15 to 20 years. References Barnes. H. F. 1937. Met hods of investigating the biometrics of thc common crane·fl y, Tipula paludosa Meigen. together with some r esults. Ann. Appl. BioI. 24:356-368. Bovien, P. 1937. Some types of association between nematodes and insects. Victensk. medd. dansk. naturh. For 101. Co penhagen, C. A. Ri etzeI. 114pp. Cohen, M and W. Steer. 1946 The co ntrol of If'a therjackets with DDT ..J. Roy. Hort. Soc. 71:130·133. Coulson, J. C. 1962. Thc biology uf Tipula subnodicornis Zetterstedt, with compara­ tive observations on Tipula paludosa Me igen. J. Anim. Ecol. 31: 1-21. Fox, C. J. S. 1957. Note on occurrence in Cape Breton Island of Tipula paludosa NI g. (Diptera:Tipulidae). Canad. Ent. 89: 288. George, K. S. 1966. A wrvey of lcalhcrjacket populations in England and Wales 1961-65. Plant Pathology 15: 1-8. r. hilarov, M. S. and L. M. Semenova. 1957. Die Kutikelpl'l'm ea bilitat bodenbewohnen­ der Tipuliden-Larven. In Zum IV . Internationalen P fl anze nschutzkongr ess Ham­ burg vom 8, bis 15 September . Z. Pflkrank, 64: 385·637. Laughlin, R. 1958. Desiccation of eggs of the crane n~ ' (Tipula oleraca, L.). Nature, Lond. 182: 613. Maerck5, H. 1939. Di e Wiesenschnaken une! ihre Bekampfung. Kranke Pflanze Hi : 107-110. Mael'cks, H. 1941. Das Sc hadauftreten del' Wiesensc hnaken (Tipuliden l in Abhan­ gigkeit von Klima. Witterung und Boden. Arb. physiol. angew. Ent. Ber!. 8: 261-275. Milne, A., R Laughlin and R. E. Coggins. 1965. The 1955 and 1959 population crashes in leatherjacket, Tipula paludosa Mei gen, in Northumberland . .1. Anim. Eco!. 34: 529-544. Morris, R. F. 1960. Newfoundland . Vegetable, field crop and other injurious insect pests predominating duri ng 1959. Canad. In s. Pest Rev. 38: 104-106. Muller-Kogler, E. 1957. Vber eine Mykose del' larven von Tipula paludosa Meig. durch Empusa sp. In Zum IV . Internationalen Pflanzenschutzkongress Hamburg vom 8, bis 15 September . Z. Pflkrankh. 64 : 385-637. Rennie, J . 1917. On the biology and economic significance of Tipula paludosa Meigen. Part I. Ann. Appl. BioI. 2: 235-240. Rennie, J . 1916. On the biological and economic significance of Tipula paludosa Meigen. II. Ann. App!. Bio!. 3: 116-137. Rennie, J . and C. H. Sutherland. 1920. On the life hi stor y of Bucentes (Siphona) geni. culata (Diptera:Tachinidae), parasite of Tipula paludosa (Diptera) and other species. Parasitology 12: 199-211. Robertson, A. G. 1939. The nocturnal activity of crane-flies (Tipulinae) as indicated by captures in a li ght trap at Rothamsted ..J. Anim. Eco!. 8:300-322. Sellke, K. 1936. Biologische und morphologische Studien an ·schadliehen Wiesen­ schnaken (Tipulidae, Dipt.) Z. wiss Zool. 148: 465-555. Sellke, K. 1937. Beobachtungen uber die Bekampfung von Wiesenschnakenlarven (Tipula paludosa Mrig, und Tipula czizeki de J.), Z. angew. Ent. 24: 277-284.