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Tipula paludosa
(European crane fly)

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Datasheet

Tipula paludosa (European crane fly)

Summary

  • Last modified
  • 16 November 2018
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Preferred Scientific Name
  • Tipula paludosa
  • Preferred Common Name
  • European crane fly
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Arthropoda
  •       Subphylum: Uniramia
  •         Class: Insecta
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    Compendia
    CAB International
    Wallingford
    Oxfordshire
    OX10 8DE
    UK
    compend@cabi.org
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Identity

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Preferred Scientific Name

  • Tipula paludosa Meigen

Preferred Common Name

  • European crane fly

Other Scientific Names

  • Tipula fimbriata Meigen 1818
  • Tipula flavolutescens Pierre 1921
  • Tipula wollastoni Lackschewitz 1936

International Common Names

  • English: crane fly, common; crane fly, European marsh; leatherjacket
  • Spanish: gusano de cuero; tipula de las huertas
  • French: tipule des prairies; tipule européenne; tipule potagere

Local Common Names

  • Denmark: mosestankelben
  • Finland: suovaaksiainen
  • Germany: Schnake, Sumpf-; Schnake, Wiesen-
  • Iran: paschehe batlagh
  • Italy: verme giacca di cuoio; zanzarone degli orti
  • Netherlands: Langpootmug
  • Norway: myrstankelbein
  • Sweden: kaerrharkrank

EPPO code

  • TIPUPA (Tipula paludosa)

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Arthropoda
  •             Subphylum: Uniramia
  •                 Class: Insecta
  •                     Order: Diptera
  •                         Family: Tipulidae
  •                             Genus: Tipula
  •                                 Species: Tipula paludosa

Notes on Taxonomy and Nomenclature

Top of page Tipula paludosa was first described by Meigen in 1830. There have been no subsequent proposed revisions and the species is considered valid. It is one of three species that comprise the T. oleracea group. It belongs to subgenus Tipula and may therefore be cited as Tipula (Tipula) paludosa Meigen.

At the beginning of the twentieth century there was confusion over the identification of the adults of the T. oleracea group. Theobald (1913) and Rennie (1916) believed that T. paludosa had two generations per year in England, UK. It is likely, however, that reports of the earlier generation were actually Tipula oleracea. In contrast, photographic evidence of T. oleracea provided by Silantjev (1931) shows female adults with short wing to body ratio, a characteristic which is typical of T. paludosa. The controversy over the identification was finally settled by De Jong (1925) who successfully separated the adults of each species and added T. czizeki De Jong to the group.

Description

Top of page Adults are large insects with females reaching 45-50 mm across. They are characterized by long, spindly legs. Antennae have 14 segments. Female wings are shorter than the abdomen. Males are generally smaller with relatively longer wings.

The number of eggs produced has been reported as between 200 and 300 (Barnes, 1937) but figures lower than this are frequently encountered in laboratory studies (RP Blackshaw, University of Plymouth, UK, personal communication, 1998). They are ovoid, shiny, black and up to 3 mm in length.

The larval body consists of 13 segments; the spiracles are found on the last segment and in the first instar small tufts of bristles can also be seen around the anal papillae. Larvae are elongated, cylindrical, a little tapered anteriorly and truncate at the rear (Coyler and Hammond, 1951). The body is enclosed in a tough integument which gives rise to the name by which they are commonly known, leatherjackets. There are four instars which vary in weight and size. The diameter of the posterior spiracles can be used to differentiate larvae and sex fourth instars (Blackshaw and Moore, 1984).

Wing length relative to abdomen length will help separate T. paludosa from the other common species found in pasture soils (see Similarities to Other Pests). However, determination of Tipulidae is a specialist task, particularly as this is the largest (known) family of Diptera (13,500 species in 300 genera). The following morphological features will help separate Tipula (Tipula) from most other subgenera and genera.

Large, wing length 13-23 mm (Coe et al., 1950). Body matt (covered in microtrichiae), not shiny, not bright coloured. Antennal flagellomeres 12 without lateral extensions; with long hairs arouns bases. Terminal segment of palpus elongate. Wing: no macrostichia in wing cells; vein Sc ending beyond origin of Rs; calypteres setose. Tarsi dark coloured.

Distribution

Top of page Much of the knowledge of the distribution of T. paludosa comes from applied research and reflects where entomologists have pursued its study. In general, it is found in temperate zones of Europe. Its spread to the east coast of North America (Jackson and Campbell, 1975) indicates a potential to colonize other areas. At present, distribution is restricted to the northern hemisphere. Suitable climatic and agronomic conditions exist elsewhere, for example in New Zealand, and further spread can be anticipated.

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Africa

Spain
-Canary IslandsPresentOosterbroek and Theowald, 1992

North America

CanadaPresentPresent based on regional distribution.
-British ColumbiaPresentCIE, 1977
-OntarioPresentPeck et al., 2006
-QuebecPresentPeck et al., 2006
USAPresentPresent based on regional distribution.
-New YorkPresentPeck et al., 2008
-OregonPresentUmble and Rao, 2004
-WashingtonPresentCIE, 1977; Jackson and Campbell, 1975

Europe

AustriaRestricted distribution****CIE, 1977
BelgiumPresentCIE, 1977; Oosterbroek and Theowald, 1992
Czechoslovakia (former)Widespread****CIE, 1977; Oosterbroek and Theowald, 1992
DenmarkPresentCIE, 1977; Oosterbroek and Theowald, 1992
Faroe IslandsPresentOosterbroek and Theowald, 1992
FinlandPresentCIE, 1977; Oosterbroek and Theowald, 1992
FranceWidespreadCIE, 1977; Ricou and Douyer, 1975; Oosterbroek and Theowald, 1992
-CorsicaPresentOosterbroek and Theowald, 1992
GermanyWidespread****CIE, 1977; Sellke, 1936; Oosterbroek and Theowald, 1992
HungaryRestricted distribution****CIE, 1977; Oosterbroek and Theowald, 1992
IrelandPresentGreenan, 1966; Oosterbroek and Theowald, 1992
ItalyPresentCIE, 1977
LuxembourgPresentOosterbroek and Theowald, 1992
NetherlandsRestricted distributionCIE, 1977; Vlug, 1990; Oosterbroek and Theowald, 1992
NorwayPresentCIE, 1977; Oosterbroek and Theowald, 1992
PolandPresentCIE, 1977; Oosterbroek and Theowald, 1992
PortugalPresentPresent based on regional distribution.
-MadeiraPresentOosterbroek and Theowald, 1992
Russian FederationRestricted distributionCIE, 1977; Oosterbroek and Theowald, 1992
-Central RussiaPresentOosterbroek and Theowald, 1992
SpainPresentOosterbroek and Theowald, 1992
SwedenWidespread****CIE, 1977; Oosterbroek and Theowald, 1992
SwitzerlandWidespreadCIE, 1977; Oosterbroek and Theowald, 1992
UKWidespread****CIE, 1977; Rennie, 1916; Oosterbroek and Theowald, 1992
-England and WalesPresentGökçe and Er, 2003
-ScotlandPresentDawson et al., 2003
Yugoslavia (former)PresentCIE, 1977; Oosterbroek and Theowald, 1992

Risk of Introduction

Top of page T. paludosa is not listed as a quarantine pest.

Habitat

Top of page T. paludosa is considered to be primarily a grassland species. Although larvae have been reported from a variety of agricultural crops, the biology and behaviour of the species determines that populations can only build up over a number of years in grassland. Other situations normally result from exposure of a crop to an existing larval population or, more rarely, direct oviposition.

Hosts/Species Affected

Top of page Adult crane flies present no concern in terms of crop damage as they do not feed on plant material. The larvae (leatherjackets), however, are recognized by farmers, advisers and gardeners as a serious threat to plant establishment (Anon., 1984). They are considered a major pest of permanent grassland, reseeds and spring cereals (French, 1969; Rayner, 1975; Newbold 1981) but are not restricted to cereal crops and grassland. Reports of leatherjacket damage from a variety of crops suggests that they are polyphagous in their feeding habits. Most of these reports, however, have not confirmed identification by, for example, rearing larvae through to adults.

Growth Stages

Top of page Seedling stage, Vegetative growing stage

Symptoms

Top of page Absolute symptoms vary depending on the crop, but there are some common features. When seedlings are attacked by larvae, typical damage is stem severance. In cereals, young plants may remain in the ground but isolated from roots. This can be superficially similar to 'frost lift' in northern climes but manifests itself as patchy damage where groups of larvae have been feeding rather than more widespread plant deaths associated with freezing soils.

In older cereal plants, leaves in contact with the ground may be severed, leaving a ragged edge.

Damage to grassland is less obvious but starts in early autumn. In the spring and early summer, extensive damage may manifest itself as dead or dying patches. Occasionally, entire swards are destroyed. The majority of damage, however, fails to yield these signs and frequently passes unnoticed.

List of Symptoms/Signs

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SignLife StagesType
Leaves / external feeding
Stems / external feeding
Whole plant / external feeding
Whole plant / frass visible

Biology and Ecology

Top of page T. paludosa has one generation per year. The adults emerge from pupation from June to September (Rennie, 1917) and peak emergence can vary regionally. Coulson (1962) found that peak emergence occurred at Moor House Nature Reserve in northern England, UK in late July and that this was 6 weeks earlier than emergence peaks obtained from Rothamsted Experimental Station further south. Blackshaw (1983a) found that peak emergence in Northern Ireland occurred mid-August to mid-September and Sellke (1936) reported that, in Germany, the peak emergence time was early September.

Female adult T. paludosa are gravid at emergence and are not able to fly far (Dobson, 1972). Mating takes place almost immediately and eggs are laid, in one batch (Cuthbertson, 1929), amongst the herbage and particularly amongst grasses. Coulson (1959) observed that female flies laid the majority of eggs close to the site of emergence. It is this behaviour on the part of adult females that causes aggregations of larvae and necessitates continuity/stability of habitat for population growth. Grassland can provide this.

The larvae hatch out after about 14 days and are a pale sandy colour. The larvae start to feed on the bases of stems and roots immediately. The larvae pass through the first two instars quickly and usually overwinter in the third-instar stage (Coulson, 1962). They can be active at temperatures as low as 5°C (Blackshaw, 1992) and so can continue to feed during periods of mild weather in winter. They become more active and feed voraciously in the spring when the soil begins to warm. When fully grown they reach approximately 3-4 cm in length and then stay within their burrows for about 6-8 weeks before making their way to the soil surface to pupate. They remain as pupae for approximately 2 weeks. In total the larvae spend a minimum of 9 months in the soil (Rennie, 1917).

Due to the impact of leatherjackets as agricultural pests, factors affecting the survival of the larvae have been well studied. The climate, and especially soil moisture, plays an important part in the survival of the larvae. The eggs and the larvae are sensitive to desiccation (Laughlin, 1958b, 1967; Meats, 1967, 1972). Milne et al. (1965) concluded that a shortage of moisture in September and October, at egg-laying time, resulted in a low larval population during the following growing season, and French (1969) also suggested a relationship between larval survival and rainfall. Blackshaw (1983b), however, found that autumn larval populations had a negative relationship with summer and autumn rainfall in Northern Ireland, and suggested that there are a number of positive and negative factors which act to sustain and suppress the population.

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Anaphes Parasite Eggs
Bacillus thuringiensis
Conidiobolus osmodes Pathogen Gökçe and Er, 2003
Entomophaga caroliniana Pathogen
Entomophaga gigantea Pathogen
Iridovirus Pathogen
Megaselia paludosa Parasite
Nucleopolyhedrosis virus Pathogen
Phaonia signata Parasite
Siphona geniculata Parasite Larvae
Tipula Irridescent Virus Pathogen Larvae

Notes on Natural Enemies

Top of page Despite the large number of natural enemies known to attack T. paludosa, there is scant evidence of significant population regulation. Generally, where feeding has been shown to occur, it coincides with the late fourth instar and prepupal phase when feeding activity is in decline.

In contrast, the parasitic wasp, Anaphes sp., was found to have attacked 44% of eggs examined in Northern Ireland (RP Blackshaw, University of Plymouth, UK, personal communication, 1998). The role of this wasp in terms of population control is not yet fully understood but it is thought that it may have future potential as a control agent (Kelly, 1989).

Given the list of predators which can feed on the larvae of Tipula spp., it must also be recognized that these larvae play an important role in the food chain of other animals, including mammals such as shrews, hedgehogs and moles. Not all scientists are in agreement with the need to reduce the population of Tipula spp. and research suggests that leatherjackets may be an important component in the diet of birds (Beintema et al., 1991; McCracken et al., 1992: Bignal et al., 1996). In some areas there may be a move to encourage leatherjacket populations to prevent the demise of other species, such as the chough (Pyrrhocorax pyrrhocorax) in Islay, in the UK (McCracken, 1990).

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Soil, sand and gravelPossible risk of moving soil. Yes

Plant Trade

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Plant parts liable to carry the pest in trade/transportPest stagesBorne internallyBorne externallyVisibility of pest or symptoms
Growing medium accompanying plants eggs; pupae Yes Pest or symptoms not visible to the naked eye but usually visible under light microscope
Roots larvae Yes Pest or symptoms usually visible to the naked eye
Plant parts not known to carry the pest in trade/transport
Bark
Bulbs/Tubers/Corms/Rhizomes
Flowers/Inflorescences/Cones/Calyx
Fruits (inc. pods)
Leaves
Seedlings/Micropropagated plants
Stems (above ground)/Shoots/Trunks/Branches
True seeds (inc. grain)
Wood

Impact

Top of page Damage occurs in both reseeds and established grassland. However, autumn reseeds with cultivations coinciding with the T. paludosa flight and oviposition period are far less vulnerable than spring reseeds, especially where the latter follows a ley. In surveys of grass reseeds in the UK (Bentley, 1984; Ellis et al., 1984; Clements et al., 1985; Bentley and Clements, 1989a, b) leatherjackets were not implicated as damaging pests in England and Wales. The majority of damage from leatherjackets is to long term leys and pastures. Blackshaw (1985) estimated that leatherjackets were responsible for in excess of £15m worth of damage in grassland in Northern Ireland alone each year. The majority of this was attributed to insidious feeding in populations below the economic threshold.

Leatherjackets have historically been associated with spring cereals and farming experience has linked extensive damage with those crops that are sown into ploughed up grass swards. Despite the long association between leatherjackets and damage to spring barley there have been few published data on yield losses. Golightly (1967) and Rayner (1969) reported experimental studies, although the similarity in reported rates of loss suggests that they may have been commenting, at least in part, on the same data.

Such rotations help determine where T. paludosa is a cereal pest. Over much of its range, the economic impact is on herbage production in grassland systems.

Given the inability of most workers to correctly identify T. paludosa larvae and the restricted dispersal ability of gravid females, doubt must be cast on many of the records of damage to other crops in the absence of corroborating evidence. The closely related species, T. oleracea, is more opportunist and may provide an alternative explanation.

Diagnosis

Top of page The problem of larval identification was tackled by Humphreys et al. (1993a) who used isoelectric focusing to distinguish between the larvae of T. paludosa and T. oleracea. Proteins were extracted from fourth-instar larvae, the subsequent staining pattern revealed protein banding patterns which consistently distinguished between the two species at all stages of development. A limitation of this method is the inability to differentiate between T. paludosa and T. czizeki. Further studies using, for example, genomic DNA technology may deliver a method that can distinguish all three species.

Detection and Inspection

Top of page T. paludosa adults can be recovered from most traps designed to sample flying insects. They have been shown, however, to favour green water traps over other colours (Blackshaw, 1983a).

Most methods for sampling larvae in grassland involve the collection of soil. For quantitative purposes this usually involves soil cores to a depth of 10-15 cm. Larvae can be recovered by heat extraction (Blasdale, 1974) or by washing the soil through a series of graduated sieves and flotation in brine (Blackshaw, 1983b). Both of these methods require laboratory facilities and most extension workers will dig up sods and hand separate soil in the field to see if larvae are present. An alternative method is to use gradual flooding of soil cores with brine as an expellent or use brine directly in the field (Stewart and Kozicki, 1987).

Since larvae tend to aggregate around plants it is possible to estimate numbers directly from along rows in drilled cereal crops. A sample of 10 x 30 cm lengths of drill across a field is considered adequate (though likely to be statistically weak). Larvae are recovered by passing the top 5 cm of soil through the fingers.

All sampling methods are generally less successful until the second larval instar has been reached.

Similarities to Other Species/Conditions

Top of page The adult stages of T. paludosa, T. oleracea and T. czizeki, although quite similar in appearance at first inspection, can nevertheless be reliably separated by several physical attributes (Den Hollander, 1975a). The distance between the eyes, the number of antennal segments and the wing length can be reliably used to distinguish between the adults (Coe et al., 1950). The wing length to body ratios are sufficiently distinct that it is reasonably easy to determine female T. oleracea (wings longer than abdomen) and T. paludosa (wings shorter than abdomen) in the field. The males of both species do not have such a marked difference in terms of wing length to body ratio but can be readily separated by the other diagnostic features.

By contrast to the adults, the larvae are morphologically very similar (Theowald, 1957, 1967; Brindle, 1959). The identification of the larvae of T. oleracea and T. paludosa centres on examination of the anal papillae as well as noting slight differences in size and colour. In practice these keys have not met with consistent success when used by researchers in the field. As a result, the life-cycle and habits of field populations are almost always accorded to T. paludosa, which is by far the most common species of the group found on the wing. Smith (1989) provided a key to larvae of species commonly found in pastures.

Eggs are more readily differentiated since T. oleracea possesses a long, coiled terminal filament.

Prevention and Control

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Early Warning Systems

Some countries, notably the UK and the Netherlands, have operated regional forecasting schemes for a number of years. These involve collecting samples from a number of fields in the winter, extracting larvae and predicting whether there will be a high or low risk year for damage. In practice, these forecasts contain little usable information for individual farmers. The oviposition behaviour of gravid females means that each field/crop still has to be sampled to determine if populations exceed the threshold. The relatively high labour requirements to achieve this means it is rarely done and so the favoured approach to this pest is reactive. There is a need to develop in-field forecasting systems that are cheap and easy to operate.

Economic Thresholds

A consensus between researchers has evolved to suggest that the economic threshold for T. paludosa larvae in grassland is about 1m/ha. There is, however, substantial evidence that populations lower than this may be worth controlling (Blackshaw, 1985) and that timing of control is important in that early insecticide applications will result in greater spring herbage yields (Newbold, 1981; Blackshaw, 1984). An added complication in calculating economic thresholds is that grass itself rarely has a value until it has been converted into an animal product so that different livestock systems result in different values.

The threshold in spring cereals is usually presented as 15 larvae from 10 x 30 cm drill lengths at 18 cm drill spacing. This equates to about 0.25 m/ha.

Thresholds do not exist for other crops but lower thresholds can be anticipated with reduced plant densities.

Chemical Control

Organo-phosphate insecticides such as fenitrothion and quinalphos are approved for use in grassland but chlorpyrifos is probably the chemical most favoured by growers and advisers (Clements et al. 1992). Leatherjackets are easily controlled in grassland by a single insecticide application but there have been few studies comparing the efficacy of approved insecticides. Newbold (1981), Blackshaw (1984) and French et al. (1990) have used chlorpyrifos as an experimental tool and reported kills in excess of 90%.

In cereals there have been even fewer published studies on chemical control of leatherjackets. Rayner (1969) reported that a 70% kill was satisfactory and later confirmed the superiority and consistency of chlorpyrifos spray treatment compared with other products (Rayner, 1975).

Cultural Control

Initial control methods used on golf courses relied on the fact that leatherjackets respond to excessive moisture in the soil by coming to the surface; greens were deliberately soaked and the larvae were trapped under a tarpaulin (Morison, 1951). In agricultural systems it has been demonstrated that although attacks are frequent on crops which follow grass in the rotation, these attacks can be prevented if the grass is ploughed in July or early August and the herbage is well buried (Anon., 1984a).

Soil cultivation is the one procedure that is known to reduce numbers of leatherjackets and markedly reduce the incidence of damage (White, 1967; Blackshaw, 1988). Mortality during the preparation of a seed-bed has been estimated to be as low as 20% (LaCroix and Newbold, 1968) but observed to be 70% in a designed experiment (Blackshaw, 1988).

Drainage has been recommended for leatherjacket control (Maercks, 1941) and this may be appropriate to areas where soil moisture availability is a limiting factor at critical stages of the life cycle, such as where Milne et al. (1965) carried out their studies.

Biological Control

Early efforts at manipulating natural enemies concentrated on Tipula Irridescent Virus (TIV) but met with little success. It has been postulated that cannibalism is the main mode of transmission of TIV (Carter, 1973a,b) and this limits the mechanism for introduction into a field population. Application of TIV in live and dead T. oleracea against field populations of leatherjackets successfully introduced the virus into the population but only at low levels (Carter, 1978a, 1978b). Kelly (1989) was able to demonstrate that TIV inoculated Galleria mellonella larvae and TIV treated bran could also be used.

The problem with using such 'packets' of TIV to try and establish epizootics is the relatively low probability of individual leatherjackets encountering a source before viability is lost. Encapsulation of the virus could increase persistence in a bait and hence may improve transmission rates. Even if this occurred, however, there still remains the problem of producing sufficient virus on a commercial scale when in vitro methods do not exist.

Bacillus thuringiensis (Waalwijk et al., 1992) and predatory nematodes (Peters and Ehlers, 1994) have been tested as control methods for leatherjackets. Significant population reductions can be achieved in the field but application costs are presently substantially higher than with conventional insecticides and may only be justified if chemicals are prohibited and there is a high risk of crop failure resulting in, for example, the need to re-establish a sward.

References

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Alford DV, 1984. A Colour Atlas of Fruit Pests. Their Recognition, Biology and Control. London, UK: Wolfe Publishing Ltd, 320 pp.

Anon., 1953. Leatherjackets, 1951-1952. Plant Pathology, 2:136.

Anon., 1981. Leatherjackets and Their Control. Publication No. 77. Edinburgh, UK: The Scottish Agricultural College.

Anon., 1989. Potato Pests. ADAS Reference Book 187, Ministry of Agriculture Fisheries and Food. London, UK: HMSO.

Barbash NM; Coll C; Dobson RM, 1991. Observations on the overwinter mortality of the larvae of Tipula paludosa Meigen Diptera: Tipulidae due to bird predation. The Scottish Naturalist, 73-84.

Beintema AJ; Thissen JB; Tensen D; Visser GH, 1991. Feeding ecology of charadriiform chicks in agricultural grasslands. Ardea, 79:31-44.

Bentley BR, 1984. Stem-boring dipterous larvae (eg Oscinella frit) as pests of direct drilled grass reseeds with particular reference to the mode of infestation and seasonality of damage inflicted on such establishments. MPhil Thesis. Reading, UK: University of Reading.

Bentley BR; Clements RO, 1989. Identity of the major pests of direct-drilled grass. Crop Protection, 8(1):49-54

Bentley BR; Clements RO, 1989. Impact of time of sowing on pest damage to direct-drilled grass and the mode of attack by dipterous stem borers. Crop Protection, 8(1):55-62

Bignal EM; McCracken DI; Stillman RA; Ovenden GN, 1996. Feeding behaviour of nesting choughs Pyrrhocorax pyrrhocorax in the Scottish Hebrides. Journal of Field Ornithology, 67:25-43.

Blackshaw RP, 1983. Some factors influencing variability in water-trap catches of Tipula spp. (Diptera: Tipulidae). Bulletin of Entomological Research, 73(4):693-699

Blackshaw RP, 1983. The annual leatherjacket survey in Northern Ireland, 1965-82, and some factors affecting populations. Plant Pathology, 32(3):345-349

Blackshaw RP, 1984. The impact of low numbers of leatherjackets on grass yield. Grass and Forage Science, 39(4):339-343

Blackshaw RP, 1985. A preliminary comparison of some management options for reducing grass losses caused by leatherjackets in Northern Ireland. Annals of Applied Biology, 107(2):279-285

Blackshaw RP, 1988. Effects of cultivations and previous cropping on leatherjacket populations in spring barley. Research and Development in Agriculture, 5(1):35-37

Blackshaw RP, 1991. Leatherjackets in grassland. Strategies for weed, disease & pest control in grassland: practical implications of recent developments and future trends. Proceedings of the British Grassland Society conference held on 27 February 1991 at Hatherley Manor Hotel, Gloucester Hurley, UK: British Grassland Society, 6.1-6.12

Blackshaw RP; Moore JP, 1984. Morphological determination of leatherjacket (Tipula paludosa) instars (Diptera: Tipulidae). Record of Agricultural Research, 32:87-89

Blasdale P, 1974. A method of turf sampling and extraction of leatherjackets. Plant Pathology, 23(1):14-16

Brindle A, 1959. Notes on the larvae of British Tipulinae Dipt., Tipulidae. Part 6, The larvae of the Tipula oleracea group. Entomologist's Monthly Magazine, 95:176-177.

Brunel E, 1973. Some problems presented by Diptera in maize crops in the west. Sciences Agronomiques Rennes, 205-207

Cameron AE, 1918. The insect association of a local environment complex in the district of Holmes Chapel, Cheshire. Transcripts of the Royal Society of Edinburgh, 52:54-78.

Campbell RL, 1975. Insecticidal control of European crane fly in Washington. Journal of Economic Entomology, 68(3):386-388

Carter JB, 1973. The mode of transmission of Tipula iridescent virus. 1. Source of infection. Journal of Invertebrate Pathology, 21(2):123-130

Carter JB, 1973a. Detection and assay of Tipula Iridescent Virus by the latex agglutination test. Journal of General Virology, 21:181-185.

Carter JB, 1974. Tipula iridescent virus infection in the developmental stages of Tipula oleracea. Journal of Invertebrate Pathology, 24(3):271-281

Carter JB, 1976. A survey of microbial, insect and nematode parasites of Tipulidae (Diptera) larvae in north-east England. Journal of Applied Ecology, 13(1):103-122

Carter JB, 1978. Field trials with Tipula iridescent virus against Tipula spp. larvae in grassland. Entomophaga, 23(2):169-174

Carter JB; Green EI; Kirkham AJ, 1983. A Tipula paludosa population with a high incidence of two pathogens. Journal of Invertebrate Pathology, 42(3):312-318

Carter JB; Griffiths C; Smith AD, 1981. A high level of parasitism by Siphona geniculata (De Geer) (Diptera: Tachinidae) in a population of Tipula paludosa (Meigen) (Diptera: Tipulidae). Entomologist's Gazette, 32(4):257-258

Chapman PA, 1994. Control of leatherjackets by natural enemies: the potential role of the ground beetle Pterostichus melanarius. Proceedings - Brighton Crop Protection Conference, Pests and Diseases, Brighton, UK, 21-24 November, 1994, vol. 2. Bracknell, UK: British Crop Protection Council, 933-934

Clements RO; Bentley BR; Moore D; Spaull AM, 1985. The impact of a range of pesticide treatments on newly sown Italian ryegrass (Lolium multiflorum) and on frit-fly (Oscinella spp.) infestation. Crop Protection, 4(2):245-254

Clements RO; Carr AJH, 1983. Pests and diseases of grass and fodder crops. In: Scopes N, Ledieu M, eds. Pest and Disease Control Handbook. Croydon, UK: British Crop Protection Council, 159-189.

Clements RO; Murray PJ; Tyas CJ, 1992. The short-term effects on wild goose behaviour of chlorpyrifos application to permanent pasture. Annals of Applied Biology, 120(1):17-23

Coe RL; Freeman P; Mattingly PF, 1950. Diptera: Nematocera Handbooks for the Identification of British Insects, Vol. 9, Part 2. London, UK: Royal Entomological Society of London.

Coll C, 1996. The development of Tipula oleracea L. (Diptera: Tipulidae as a pest of winter cereals: the role of oilseed rape (Brassica napus L.). PhD Thesis. Aberdeen, UK: University of Aberdeen.

Coulson JC, 1959. Observations on the Tipulidae Diptera. of the Moor House Nature Reserve, Westermorland. Transactions of the Royal Entomology Society of London, 111:157-174.

Coulson JC, 1962. The biology of Tipula subnodicornis Zetterstedt with comparative observations on Tipula paludosa Meigen. Journal of Animal Ecology, 31:1-21.

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GISD/IASPMR: Invasive Alien Species Pathway Management Resource and DAISIE European Invasive Alien Species Gatewayhttps://doi.org/10.5061/dryad.m93f6Data source for updated system data added to species habitat list.

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