Liriomyza sativae (vegetable leaf miner)
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- Risk of Introduction
- Hosts/Species Affected
- Host Plants and Other Plants Affected
- Growth Stages
- List of Symptoms/Signs
- Species Vectored
- Biology and Ecology
- Natural enemies
- Notes on Natural Enemies
- Pathway Vectors
- Plant Trade
- Detection and Inspection
- Similarities to Other Species/Conditions
- Prevention and Control
- Links to Websites
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Liriomyza sativae Blanchard, 1938
Preferred Common Name
- vegetable leaf miner
Other Scientific Names
- Agromyza subpusilla
- Lemurimyza lycopersicae Pla & de la Cruz, 1981
- Liriomyza canomarginis Frick, 1952
- Liriomyza guytona Freeman, 1958
- Liriomyza minutiseta Frick, 1952
- Liriomyza munda Frick, 1957
- Liriomyza propepusilla Frost, 1954
- Liriomyza pullata Frick, 1952
- Liriomyza subpusilla
- Liriomyza verbenicola Hering, 1951
International Common Names
- English: cabbage leaf miner; leaf miner of vegetables; melon leaf miner; serpentine vegetable leaf miner
- Spanish: minador de la higuerilla; minador de las hojas de col; minador del tomate; minador serpentina de las hortalizas
- French: mineuse maraichère
Local Common Names
- UK: South American leaf miner
- USA: tomato leaf miner
- LIRISA (Liriomyza sativae)
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Arthropoda
- Subphylum: Uniramia
- Class: Insecta
- Order: Diptera
- Family: Agromyzidae
- Genus: Liriomyza
- Species: Liriomyza sativae
Notes on Taxonomy and NomenclatureTop of page Liriomyza sativae was originally described in Argentina from material bred from the leaves of Medicago sativa. Since then it has been found in North, Central and South America, the Caribbean Islands, Africa, some Pacific Islands and some Asian countries. It has been introduced into Europe, usually via imports for glasshouse cultivation. Its many synonyms are evidence of its importance as a widespread pest over a large area (Spencer, 1973a; 1981, 1986).
L. sativae is one of a group of six polyphagous Liriomyza flies which have developed into serious agricultural pests in all parts of the world. They may be divided into two sub-groups: L. huidobrensis, L. trifolii, L. sativae and L. brassicae, originally discovered in the Americas, forming one group, and L. bryoniae and L. strigata from Europe forming the other.
Spencer (1973a; 1990) details a considerable increase in the number of species identified, no doubt in part due to increased work done on the Agromyzidae, but also to increased international commerce in fresh plant material creating opportunities for the establishment of flies across continents.
The extent of the darkening of the hind-margin of the eye and of the mesopleura [anepisternum] is not constant and the attachment of too much importance to these characters led Frick (1952) incorrectly to describe L. sativae as three species in Hawaii as L. pullata, L. canomarginis and L. minutiseta, thus emphasizing the need for caution in the use of external morphology alone. However, the hind margin of the eye is always darker than in L. trifolii and the mesopleura is always paler than in L. huidobrensis (Spencer, 1973a). Steyskal (1964) synonymized L. guytona with L. munda. Spencer (1973a) then synonymized L. munda, L. pullata, L. canomarginis and L. minutiseta with L. sativae.
DescriptionTop of page Descriptions of L. sativae refer to fresh materials. Dry specimens may be distorted due to the manner in which they have been preserved. Also, the age of the specimen, when killed, will have some effect on its preservation characteristics.
L. sativae eggs are 0.2-0.3 x 0.1-0.15 mm, off-white and slightly translucent.
This is a legless maggot with no separate head capsule, transparent when newly hatched but colouring up to a yellow-orange in later instars, up to 3 mm long. Both larvae and puparia have a pair of posterior spiracles terminating in three cone-like appendages. Spencer (1973a) describes distinguishing features of the larvae. Petitt (1990) describes a method of identifying the different instars of the larvae.
Oval, slightly flattened ventrally, 1.3-2.3 x 0.5-0.75 mm with variable colour, pale yellow-orange, darkening to golden-brown. The pupa has posterior spiracles on a pronounced conical projection, each with three distinct bulbs, two of which are elongate. Pupariation occurs outside the leaf, either on the leaf or on the soil beneath the leaf.
Menken and Ulenberg (1986) describe a method of distinguishing L. sativae from L. bryoniae, L. huidobrensis, and L. trifolii using allozyme variation patterns as revealed by gel electrophoresis.
L. sativae is very small (1-1.3 mm body length, up to 1.7 mm in female with wings 1.3-1.7 mm.) The mesonotum is shiny black to the edge of a bright yellow scutellum; the face, frons and third antennal segment are bright yellow. Males and females are generally similar in appearance.
L. sativae are not very active fliers, and in crops showing active mining many flies may be seen walking rapidly over the leaves with only short jerky flights to adjacent leaves.
Head: the frons projects very slightly above the eye, just less than 1.5 times the width of the eye. There are two equal ors and two ori (the lower one weaker). Orbital setulae are sparse and reclinate. The jowls are deep (almost 0.33 times height of the eye at the rear); the cheeks form a distinct ring below the eye. The third antennal segment is small, round and noticeably pubescent, but not excessively so.
Mesonotum: 3+1 dc (3 dorsocentral bristles after the dorsal suture and 1 before it, nearer the head end), the third and fourth are substantially weaker. The distance between the first and second is up to twice that between second and third. The second, third and fourth are almost equidistant; acr appear irregularly in four rows.
Wing: length 1.3-1.7 mm, the discal cell is small. The last section is M<(sub)3+4> from 3-4 times the length of the penultimate one.
Genitalia: the shape of the distiphallus is fairly distinctive but could be misidentified for L. trifolii. Identification using the male genitalia should only be undertaken by specialists.
Colour: the head (including the antenna and face), is bright yellow. The hind margin of the eye is black, vte usually on black ground, the dark colour diminishes towards the upper orbits and becomes paler, brownish towards the base of vti, which may be just on dark ground or on the yellow. The mesopleura is predominantly yellow, with a variable dark area, from a slim grey bar along the base to extensive darkening reaching higher up the front margin than the back margin. The sternopleura is largely filled by a black triangle, but always with bright yellow above. The femora and coxa are bright yellow. The tibia and tarsi are darker. The fore-legs are brownish-yellow and the hind legs are brownish-black. The abdomen is largely black but the tergites are variably yellow, particularly at the sides. The squamae are yellowish, with a dark margin and fringe. Although individual specimens may vary considerably in colour, the basic pattern is consistent.
DistributionTop of page It is difficult to give accurate distributional notes on L. sativae at present, as there is every evidence that the fly is rapidly expanding its presence and colonizing most habitats to which it is introduced. An example is its present status in China, where it is widespread (Institute of Zoology, Beijing, China; report in preparation). Originally recognized as present in Sanya, Hainan Provinces in 1993 ( Xie-Qonh Hua et. al., 1997), it has quickly spread north and west to most Provinces since that time, causing serious damage in some areas. This is probably the true situation in most countries where it has been introduced. It will take some years before a more settled picture can be given, where a combination of natural climatic restrictions and man's effort at eradication will stabilize the flies' progress.
L. sativae has been identified as an A1 risk in the Netherlands (OEPP/EPPO, 1984) and the UK (EPPO, 1984).
See also CABI/EPPO (1998, No. 95).
Distribution TableTop of page
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/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Bangladesh||Present||Santosh and Bhuiya, 2014|
|Cambodia||Absent, unreliable record||EPPO, 2014|
|China||Restricted distribution||EPPO, 2014|
|-Fujian||Present||CABI/EPPO, 2006; EPPO, 2014|
|-Hubei||Present||Li et al., 2006|
|-Shandong||Present||Zeng et al., 2002|
|-Shanghai||Present||Yuan et al., 2000|
|-Shanxi||Present||CABI/EPPO, 2006; EPPO, 2014|
|-Sichuan||Present||CABI/EPPO, 2006; EPPO, 2014|
|-Xinjiang||Present||Zhu et al., 2004|
|India||Restricted distribution||EPPO, 2014|
|-Uttar Pradesh||Present||EPPO, 2014|
|-Java||Present||CABI/EPPO, 2006; EPPO, 2014|
|Iran||Present||CABI/EPPO, 2006; EPPO, 2014|
|Japan||Restricted distribution||EPPO, 2014|
|-Honshu||Restricted distribution||EPPO, 2014|
|-Kyushu||Restricted distribution||EPPO, 2014|
|-Ryukyu Archipelago||Restricted distribution||EPPO, 2014|
|Laos||Absent, unreliable record||EPPO, 2014|
|Malaysia||Present||CABI/EPPO, 2006; EPPO, 2014|
|-Peninsular Malaysia||Present||CABI/EPPO, 2006; EPPO, 2014|
|Oman||Present||1990||Deeming, 1992; EPPO, 2014|
|Pakistan||Absent, unreliable record||EPPO, 2014|
|Sri Lanka||Present||CABI/EPPO, 2006; EPPO, 2014|
|Thailand||Restricted distribution||1994||Martinez, 1994; EPPO, 2014|
|Turkey||Restricted distribution||EPPO, 2014|
|Uzbekistan||Restricted distribution||EPPO, 2014|
|Yemen||Present, few occurrences||Deeming, 1992; EPPO, 2014|
|Cameroon||Present||Martinez and Bordat, 1996; EPPO, 2014|
|Congo||Absent, unreliable record||EPPO, 2014|
|Ethiopia||Absent, unreliable record||EPPO, 2014|
|South Africa||Absent, unreliable record||EPPO, 2014|
|Sudan||Present||Martinez and Bordat, 1996; EPPO, 2014|
|Tanzania||Absent, unreliable record||EPPO, 2014|
|Zimbabwe||Restricted distribution||EPPO, 2014|
|Canada||Restricted distribution||EPPO, 2014|
|-Ontario||Present||McClanahan, 1980; EPPO, 2014|
|Mexico||Present||EPPO, 2014; Medina et al., 2014|
|USA||Restricted distribution||EPPO, 2014|
|-Alabama||Present||Spencer and, 1986; EPPO, 2014|
|-Arizona||Present||EPPO, 1990; EPPO, 2014|
|-California||Present||Spencer and, 1986; EPPO, 1990; EPPO, 2014|
|-Florida||Present||Spencer and, 1986; EPPO, 1990; EPPO, 2014|
|-Georgia||Present||Chalfant and Young, 1984; EPPO, 1990; EPPO, 2014|
|-Hawaii||Present||Spencer and, 1986; EPPO, 1990; EPPO, 2014|
|-Indiana||Present||York, 1988; EPPO, 2014|
|-New Jersey||Present||EPPO, 2014|
|-Ohio||Present||Spencer and, 1986; EPPO, 1990; EPPO, 2014|
|-South Carolina||Present||Spencer and, 1986; EPPO, 2014|
|-Texas||Present||Spencer and, 1986; EPPO, 2014|
Central America and Caribbean
|Antigua and Barbuda||Present||EPPO, 2014|
|Bahamas||Restricted distribution||EPPO, 2014|
|Barbados||Restricted distribution||Spencer and, 1986; EPPO, 2014|
|Costa Rica||Present||EPPO, 2014|
|Dominican Republic||Present||EPPO, 2014|
|Jamaica||Restricted distribution||Spencer and, 1986; EPPO, 2014|
|Netherlands Antilles||Present||EPPO, 2014|
|Nicaragua||Present||Rosset et al., 1987; EPPO, 2014|
|Puerto Rico||Present||EPPO, 2014|
|Saint Kitts and Nevis||Present||EPPO, 2014|
|Saint Lucia||Present||Introduced||Invasive||EPPO, 2014|
|Saint Vincent and the Grenadines||Widespread||EPPO, 2014|
|Trinidad and Tobago||Present||EPPO, 2014|
|Argentina||Widespread||Spencer and, 1986; EPPO, 2014|
|Brazil||Restricted distribution||Spencer and, 1986; EPPO, 2014|
|-Parana||Present||Lorini and Foerster, 1987; EPPO, 2014|
|-Rio de Janeiro||Present||EPPO, 2014|
|-Rio Grande do Norte||Present||Spencer, 1990; EPPO, 2014|
|Chile||Restricted distribution||Spencer and, 1986; EPPO, 2014|
|Colombia||Restricted distribution||EPPO, 2014|
|French Guiana||Present||EPPO, 2014|
|Peru||Restricted distribution||Spencer and, 1986; EPPO, 2014|
|Suriname||Absent, unreliable record||EPPO, 2014|
|Venezuela||Restricted distribution||Spencer, 1973b; Spencer and, 1986; EPPO, 2014|
|Croatia||Absent, confirmed by survey||EPPO, 2014|
|Estonia||Absent, confirmed by survey||EPPO, 2014|
|Finland||Absent, intercepted only||EPPO, 2014|
|Netherlands||Absent, intercepted only||EPPO, 2014||Absent, intercepted only, confirmed by survey.|
|Poland||Absent, invalid record||CABI/EPPO, 2006; EPPO, 2014|
|UK||Absent, intercepted only||Ledieu and Bartlett, 1983; EPPO, 1984; EPPO, 2014|
|American Samoa||Widespread||EPPO, 2014|
|Australia||Absent, formerly present||IPPC, 2008; EPPO, 2014; IPPC, 2015|
|-Queensland||Present||EPPO, 2014; Blacket et al., 2015; IPPC, 2015; IPPC, 2017|
|Cook Islands||Restricted distribution||EPPO, 2014|
|French Polynesia||Present||EPPO, 2014|
|Guam||Restricted distribution||Johnson, 1993; EPPO, 2014|
|Micronesia, Federated states of||Present||Johnson, 1993; EPPO, 2014|
|New Caledonia||Restricted distribution||EPPO, 2014|
|Northern Mariana Islands||Present||EPPO, 2014|
|Papua New Guinea||Present||Blacket et al., 2015|
Risk of IntroductionTop of page L. sativae has become a major pest of a wide variety of ornamental and vegetable crops, most notably tomatoes, chrysanthemums and celery, particularly under glass. Its importance has grown rapidly in the Americas since the 1970s, and distribution through commerce to other parts of the world is now producing serious problems everywhere, particularly in warmer climates. L. sativae is therefore listed as an A1 quarantine pest by EPPO (1984).
HabitatTop of page L. sativae will spread in any suitable habitat with a warm temperature, especially in temperature-controlled glasshouses where reproduction can become almost continuous. Particularly suitable are monocultures of primary hosts (such as Solanaceae, Fabaceae and Asteraceae) which can give rise to very large fluctuations in populations, particularly in glasshouse conditions where natural migration of hymenopterous parasites is difficult. At harvest, when the primary host is removed, there are usually enough suitable wild hosts locally to act as a reservoir and ensure continuation of L. sativae until the fields, or glasshouses, are again planted with preferred hosts.
Hosts/Species AffectedTop of page L. sativae is able to colonize a wide range of plants (primarily although not exclusively Solanaceae, Fabaceae and Asteraceae), thus ensuring success in establishing itself at any opportunity (see Spencer, 1990).
Host Plants and Other Plants AffectedTop of page
|Abelmoschus esculentus (okra)||Malvaceae||Main|
|Apium graveolens (celery)||Apiaceae||Main|
|Arachis hypogaea (groundnut)||Fabaceae||Main|
|Beta vulgaris var. saccharifera (sugarbeet)||Chenopodiaceae||Main|
|Brassica oleracea (cabbages, cauliflowers)||Brassicaceae||Main|
|Brassica rapa cultivar group Mizuna||Brassicaceae||Main|
|Brassica rapa subsp. rapa (turnip)||Brassicaceae||Main|
|Brassicaceae (cruciferous crops)||Brassicaceae||Main|
|Cajanus cajan (pigeon pea)||Fabaceae||Main|
|Capsicum annuum (bell pepper)||Solanaceae||Main|
|Cicer arietinum (chickpea)||Fabaceae||Other|
|Citrullus lanatus (watermelon)||Cucurbitaceae||Main|
|Cucumis melo (melon)||Cucurbitaceae||Main|
|Cucumis sativus (cucumber)||Cucurbitaceae||Main|
|Cucurbita maxima (giant pumpkin)||Cucurbitaceae||Main|
|Cucurbita pepo (marrow)||Cucurbitaceae||Main|
|Dahlia pinnata (garden dahlia)||Asteraceae||Other|
|Daucus carota (carrot)||Apiaceae||Main|
|Fabaceae (leguminous plants)||Fabaceae||Main|
|Lactuca sativa (lettuce)||Asteraceae||Main|
|Lathyrus odoratus (sweet pea)||Fabaceae||Main|
|Medicago sativa (lucerne)||Fabaceae||Main|
|Nicotiana tabacum (tobacco)||Solanaceae||Main|
|Ocimum basilicum (basil)||Lamiaceae||Main|
|Phaseolus vulgaris (common bean)||Fabaceae||Main|
|Pisum sativum (pea)||Fabaceae||Main|
|Raphanus sativus (radish)||Brassicaceae||Main|
|Ricinus communis (castor bean)||Euphorbiaceae||Other|
|Solanum lycopersicum (tomato)||Solanaceae||Main|
|Solanum melongena (aubergine)||Solanaceae||Main|
|Solanum tuberosum (potato)||Solanaceae||Main|
|Spinacia oleracea (spinach)||Chenopodiaceae||Main|
|Zea mays (maize)||Poaceae||Main|
Growth StagesTop of page Flowering stage, Fruiting stage, Seedling stage, Vegetative growing stage
SymptomsTop of page L. sativae feeding punctures on the upper side of the leaves appear as white speckles between 0.13 and 0.15 mm in diameter. Oviposition punctures are usually smaller (0.05 mm) and are usually more uniformly round.
Mines are usually white with dampened black and dried brown areas. They are typically serpentine, tightly coiled, or of irregular shape, increasing in width as larvae mature. The frass is distinctive in being deposited in black strips alternately at either side of the mine (Spencer, 1973a). In larger leaves, the mines often form an irregular 'U' shape.
Fungal destruction of the leaf may also occur as a result of infection introduced from other sources during breeding activity. Wilt may occur, especially in seedlings.
List of Symptoms/SignsTop of page
|Leaves / abnormal colours|
|Leaves / abnormal leaf fall|
|Leaves / external feeding|
|Leaves / internal feeding|
|Leaves / necrotic areas|
|Leaves / rot|
|Leaves / wilting|
Biology and EcologyTop of page Egg
L. sativae eggs are inserted just below the leaf surface. Eggs hatch in 2-5 days according to temperature. Harris and Tate (1933) give 4-7 days at 24°C. Many eggs may be laid on a single leaf.
The duration of L. sativae larval development also depends on temperature and probably on host plant. Stegmaier (1966) reports up to 80 larvae per leaf in Ricinus. Up to 24 generations can occur during the year (Xie-QongHua et al., 1997; Chen-ZaiLiao et al., 1998; Zeng-Ling et al., 1998) although 10-14 is more normal, breeding probably only being restricted by the availability of fresh plant growth in suitable hosts (Spencer, 1973).
Puparial development will vary according to season and temperature. Adult emergence occurs 7-14 days after pupariation at temperatures between 20 and 30°C (Leibee, 1982). Wolfenbarger (1947) gives 24-28 days for the complete cycle, in Florida (USA) during December-January (winter period).
Peak emergence of L. sativae adults occurs before midday (McGregor, 1914). Males usually emerge before females. Mating takes place from 24 hours after emergence and a single mating is sufficient to fertilize all eggs laid.
Female flies puncture the leaves of the host plants causing wounds which serve as sites for feeding or oviposition. Feeding punctures cause the destruction of a large number of cells on the host plant and are more clearly visible to the naked eye. About 15% of oviposition punctures made by L. sativae contain viable eggs (Parrella et al., 1981). Males are unable to puncture the leaves but have been observed feeding at punctures made by females. Both males and females feed on dilute honey (in the laboratory) and take nectar from flowers (EPPO, 1990).
Both male and female L. sativae may act as vectors for disease by transference during feeding or egg laying, but are not inherent carriers of disease.
Natural enemiesTop of page
|Natural enemy||Type||Life stages||Specificity||References||Biological control in||Biological control on|
|Bacillus thuringiensis kurstaki||Pathogen|
|Bacillus thuringiensis thuringiensis||Pathogen|
|Closterocerus utahensis||Parasite||Larvae||USA; Hawaii||watermelons|
|Cothonaspis pacifica||Parasite||USA; Hawaii||watermelons|
|Diglyphus begini||Parasite||Larvae||Hawaii; USA; Hawaii||chrysanthemums; watermelons|
|Ganaspidium hunteri||Parasite||Larvae||USA; Hawaii||watermelons|
|Gronotoma adachiae||Parasite||Abe and Konishi, 2012|
|Halticoptera circulus||Parasite||Larvae||USA; Hawaii||watermelons|
|Opius dimidiatus||Parasite||Hawaii; USA||chrysanthemums|
|Opius dissitus||Parasite||Eggs/Larvae||USA; Hawaii||watermelons|
|Zaeucoila unicarinata||Parasite||Brazil; Parana||cucumbers|
Notes on Natural EnemiesTop of page
The families Eulophidae and Braconidae and near relatives (very small hymenopterous parasitic wasps), generally termed the Parasitica, appear to be the natural enemies of L. sativae. There is a worldwide distribution of these insects and in natural conditions L. sativae, like all Agromyzidae, is kept to low, balanced populations by their efforts. Waterhouse and Norris (1987) have compiled a complete list of recorded natural enemies of L. sativae.
However, since 1950, with the rise in the use of largely oil-based insecticides, many of the natural predators of L. sativae have been shown to be more susceptible to the insecticides than L. sativae itself. Consequently the survival of resistant strains of Agromyzidae have had no enemies to control them, resulting in very large and damaging populations.
Since the 1970s (Spencer, 1973a) damage done by artificial chemical control has been recognized and biological controls are now being studied and implemented to provide safer and more effective population controls.
Commercial breeding is now a regular method of producing early supplies of these parasites, particularly for glasshouse crops, thus reducing the initial population growth of L. sativae (McClanahan, 1980; Johnson, 1993). See also McClanahan (1980), Johnson (1993), Cruz and Cardona (1998), Xu-ZaiFu et al. (1998).
Pathway VectorsTop of page
Plant TradeTop of page
|Plant parts liable to carry the pest in trade/transport||Pest stages||Borne internally||Borne externally||Visibility of pest or symptoms|
|Fruits (inc. pods)||eggs; larvae; pupae||Yes||Pest or symptoms usually visible to the naked eye|
|Leaves||eggs; larvae; pupae||Yes||Pest or symptoms usually visible to the naked eye|
|Seedlings/Micropropagated plants||eggs; larvae; pupae||Yes||Pest or symptoms usually visible to the naked eye|
|Plant parts not known to carry the pest in trade/transport|
|Growing medium accompanying plants|
|Stems (above ground)/Shoots/Trunks/Branches|
|True seeds (inc. grain)|
ImpactTop of page L. sativae is the most serious of the agromyzid pests, causing severe damage and loss of yield in many of the southern states of America, and also in South America. Its spread to other areas has become a matter of considerable concern. Losses of 80% have been reported for celery in Florida; tomato yields in many parts of its range and many other crops may be seriously reduced by its activities. Medicago sativa in Argentina has been seriously damaged, with estimates of up to 80% losses (Spencer, 1973a). Young plants are particularly susceptible to damage and consequent reduced efficiency or death, whilst older plants may also be seriously damaged through leaf loss due to many mines occurring in each leaf.
In most of its habitats L. sativae presents a serious threat as a pest of many agricultural crops (particularly Solanaceae and Fabaceae, but has also been recorded on seven other families). It is difficult to eradicate because of its ability to survive in many weed plants which normally occur in areas adjacent to crop fields.
Detection and InspectionTop of page The spread of Liriomyza species throughout international commerce means that identification of individual infestations must be done by a specialist of the family (Spencer, 1973a).
L. sativae flies may be detected flying closely around host plants or moving erratically and rapidly upon the leaf surfaces. Inspection of the leaf surface will reveal punctures of the epidermis and the obvious whitish mines with linear grains of frass at intervals along the length of the mine.
For accurate identification, examination of the leaf mine and all stages of development are crucial.
Successful L. sativae larvae will be found still feeding at the end of the mine, or the mine will end with a small convex slit in the epidermis. Sometimes the puparia, or its exuviae, may be found adhering to the leaf surface, although in most cases the fully fed larva will have found its way to the ground beneath the plant to pupariate. This is especially true in hot dry conditions where the larva/puparia would quickly desiccate if exposed on the leaf surface. Empty puparial cases are split at the anterior end, but not usually separated from the rest of the case.
Mined leaves should be collected into polythene bags and transferred to a press as soon as possible. Leaves containing larvae intended for breeding should be collected into individual polythene bags, which on return to the laboratory should be slightly over-pressurized by blowing into them before sealing the end. Blowing up the bag by mouth and sealing it adds valuable carbon dioxide to the moist air mix. Constant attention is required to ensure that pupae are transferred to individual tubes until the fly emerges. If the plant material begins rotting, good material with feeding larvae must be removed to more sanitary conditions.
When pupae are observed they can be removed to tubes containing a layer of fine sand or a small strip of blotting or filter paper. This should be kept damp, never wet, until the imago emerges.
On emergence, the fly should be kept for at least 24 hours to harden up. Do not allow condensation to come into contact with the fly, or it will stick to the water film and be damaged.
Collection of the adult L. sativae is done by netting. Sticky traps, especially yellow ones, placed near host plants provide a very effective method of collection and estimation of infestation. For positive identification, leaves containing larvae or puparia may be picked and kept in a humid, not wet, atmosphere in a sealed plastic bag. If the puparial stage is collected, these may be separated from the leaf. Great care is needed to ensure that no damage is done to the puparial skin or death will almost certainly follow. The pupariae may be stored in glass tubes on a layer of clean sand, or better still, thick filter paper. The tube must have an atmosphere of very high humidity, but without condensation. When the fly emerges, it should be allowed to harden for 24 hours before killing for identification purposes, but again care should be taken to ensure that there is no condensation present in the tube.
Newly emerged adult L.sativae are generally softer than specimens aged for several days and may crinkle as drying proceeds, especially at the head. The ptilinal sac may still protrude from the suture between the frons and face, obliterating some important characteristics. Adults should be dried slowly in the dark in a sealed receptacle over blotting paper. If preserving wet is preferred, the live specimen should be dropped into 20-40% alcohol, and transferred to 70-90% alcohol after 2 days.
Similarities to Other Species/ConditionsTop of page Liriomyza species, in general, may be recognized by their black (sometimes brilliantly black) and yellow colouring. Particularly, the scutellum is usually bright yellow and distinctive. All Liriomyza species are similar and may be mistaken for each other on quick examination. These are L. sativae (origins probably in South America); L. huidobrensis (origins in South America); L. trifolii (origins probably the Caribbean/Florida); L. bryoniae (origins in Europe); L. strigata (origins in Europe); L. congesta (origins in Europe/Western Asia); L. brassicae (origins probably North America).
L. sativae may often be distinguished from other species by the brilliant, shiny, black mesonotum, although the paler forms of L. brassicae may be confused with it. In particular, the scutellum of L. sativae is usually yellow and distinctive. In such cases final identification must be made from a study of the male genitalia.
The spread of Liriomyza species through international commerce and the similarities between the seven species means that identification of individual infestations must be done by specialists (Spencer, 1973).
Menken and Ulenberg (1986) describe a method of distinguishing L. sativae from L. bryoniae, L. huidobrensis and L. trifolii, using allozyme variation patterns as revealed by gel electrophoresis.
Prevention and ControlTop of page
The release of gamma-irradiated and hence sterile male L. sativae or Parasitica in glasshouses has been tested with considerable success. Süss et al. (1986) used L. trifolii on chrysanthemums, but the technique may be applied irrespective of species.
Parasitica release has also proved a useful technique in the control of L. sativae. Wharton (1984) used Opius dissectus reared from L. sativae on sorghum, tomatoes and Phaseolus spp. for biological control experiments.
Since 1950, with the rise in the use of largely oil-based insecticides, and chlorinated chemicals, many of the natural predators of L. sativae have been shown to be more susceptible to the insecticides than L. sativae itself. Consequently the survival of resistant strains (such as Agromyzidae) have had no enemies to control them, resulting in very large and damaging populations.
Regho-Filho et al. (1993) have reported on preventive spraying of seedlings with residual insecticides such as abermectin or pyrethrum sprays to reduce early egg laying. Oliveira et al. (1991) have used the insecticides trichlorfon and diazinon in an integrated control programme, but found that although the number of leaflets mined was reduced, permethrin could significantly improve the yield of tomatoes, and the presence of a natural enemy of the genus Opius, which attacks the puparial stage, was an important regulator. Within this study, a large number of tomato cultivars were tested for resistance to attack by L. sativae.
Limited success has also been obtained by exposing yellow sticky traps close above the growing plants, as the males tend to fly low between plants and in general are more mobile than the females. This practice helps to reduce the number of sexually active males, in particular, and so reduce mating probability (Espino et al., 1988; Liu et al., 1992).
Bordat et al. (1988) made studies of the effect of entomogenous fungal strains on L. sativae pupae and found great reductions (sometimes as much as 80%) in the emergence of adults.
General hygiene conditions in horticulture and farming should be maintained by clearing all weeds and treating glasshouse soils where practical, to destroy pupae. Weeds and waste plant material, if infested, should be burned and not composted (Velez et al., 1980).
All stages are killed within a few weeks by cold storage at 0°C. Newly laid eggs are the most resistant stage and it is recommended that cuttings of infested ornamental plants (such as carnations and chrysanthemums) be maintained under normal glasshouse conditions for 3-4 days after lifting to allow eggs to hatch. Subsequent storage of plants at 0°C for 1-2 weeks should then kill off the larvae of leaf miner species (Webb and Smith, 1970).
To avoid the introduction of L. sativae (and other leaf miner species L. huidobrensis and Amauromyza maculosa [Nemorimyza maculosa]), EPPO (OEPP/EPPO, 1990) recommends that propagating material (except seeds) of Capsicum, carnations, celery, chrysanthemums, Cucumis, Gerbera, Gypsophila, lettuces, Senecio hybridus and tomatoes from countries where L. sativae occurs must have been inspected at least monthly during the previous 3 months and found to be free from L. sativae.
A phytosanitary certificate should be required for cut flowers and for harvested and transported vegetables with leaves.
ReferencesTop of page
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Distribution MapsTop of page
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