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(Coleoptera, Diptera, Hymenoptera and Lepidoptera)
by
Brian Pitkin, Willem Ellis, Colin Plant and Rob Edmunds
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TRIFOLIUM.
Clovers and Hop Trefoil. [Fabaceae]
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Seventy
species of Trifolium are recorded in Britain. These include
Hop Trefoil (T. campestre), Strawberry Clover (T. frageriferum),
Alsike Clover (T. hybridum), Crimson Clover (T. incarnatum),
Sulphur Clover (T. ochroleucon), Hungarian Clover (T.
pannonicum), Red Clover (T. pratense) and White Clover
(T. repens). The BSBI provide a downloadable plant crib for vegetative widespread species of Trifolium.
Sixteen British miners are recorded on Trifolium.
A key to the European miners recorded on Trifolium is provided in Bladmineerders van Europa. |
 White
clover
Trifolium repens
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Key for the identification of the known mines of British
Diptera recorded on Trifolium
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Note: Diptera larvae may live in a corridor mine, a corridor-blotch mine, or a blotch mine, but never in a case, a rolled or folded leaf, a tentiform mine or sandwiched between two more or less circular leaf sections in later instars. Pupation never in a cocoon. All mining Diptera larvae are leg-less maggots without a head capsule (see examples). They never have thoracic or abdominal legs. They do not have chewing mouthparts, although they do have a characteristic cephalo-pharyngeal skeleton (see examples), usually visible internally through the body wall. The larvae lie on their sides within the mine and use their pick-like mouthparts to feed on plant tissue. In some corridor miners frass may lie in two rows on alternate sides of the mine. In order to vacate the mine the fully grown larva cuts an exit slit, which is usually semi-circular (see Liriomyza huidobrensis video). The pupa is formed within the hardened last larval skin or puparium and as a result sheaths enclosing head appendages, wings and legs are not visible externally (see examples).
See Key to non-Diptera.
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1a > Leaf-miner: Linear mine in sepals and leaves. Puparium orange-brown |
On Anthyllis, but not yet on Trifoloium,in Britain and this and other Fabaceae elsewhere.
Known only from Devon in Britain and continental Europe.
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Phytomyza
brischkei Hendel, 1922 [Diptera: Agromyzidae]. |
1b > Leaf-miner: A distinctive mine primarily above mid-rib, with irregular short
lateral offshoots into leaf blade. Pupation external (Spencer, 1972:
51 (fig. 172), 55; Spencer, 1976:
270, 271 (fig. 486)).
Branched,
whitish, upper-surface corridor; main axis overlying the midrib;
side branches overlying the main lateral veins. (In Campanula and Phyteuma the mine is much less branched, sometimes nothing
more than a corridor on top of the midrib). Frass in rather long
strings. Usually the mines begins as a long and narrow, shallow,
tortuous lower-surface corridor that ends upon the midrib but otherwise
is not associated with the leaf venation. Often this initial corridor
is filled with callus, and then even less conspicuous. Pupation
outside the mine.
A
linear mine on the upper surface, usually following the midrib and
showing side branches along the veins. The frass is in strings. |
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Polyphagous. On more than 40 host genera in 15 families, but not yet on Trifolium, in Britain,. Widespread
throughout Britain. Also recorded in the Republic of Ireland.
Widespread in continental Europe.
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Liriomyza strigata (Meigen, 1830) [Diptera: Agromyzidae]. |
1c > Leaf-miner: Initially
a linear mine running towards apex of leaf which then turns back
and develops into a blotch in the area of the mid-rib (Spencer, 1972b: 37, fig. 114A; Spencer,
1976: 110, fig. 177).
Upper-surface
corridor along the leaf margin, running from the leaf base to the
tip. Once at the tip the corridor goes down over the midrib, widening
considerably. Frass in the corridor part in a broad green central
band, with minute black particles at either side. In the final,
blotchy part frass in coarse black grains that may sometimes stick
together. Pupation outside the mine.
Puparium reddish brown. |

Mines of Agromyza frontella on Medicago sativa
Image: © Whitney Cranshaw (Bladmineerders van Europa) |
On Medicago, Melilotus and Trifolium in Britain and elsewhere. Widespread in Britain
and continental Europe.
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Agromyza
frontella (Rondani, 1875) [Diptera: Agromyzidae]. |
1d > Leaf-miner: Larva
producing a short narrow linear mine not associated with the margin,
which then develops into a conspicuous white blotch centred on the
mid-rib. At the end of feeding the larva lies for one or two days
along the mid-rib. Pupation external (Spencer, 1972b: 37 (fig. 114b); Spencer,
1976: 126, fig. 213A).
Broad
and short hook-like upper-surface corridor, ending in a little blotch
in the centre of a leaflet. The blotch is upper-surface as well,
but has some deep spots, giving the mine a mottled appearance when
hold against the light. Frass in the blotch in a few large lumps.
Pupation outside the mine.
A
short, broad gallery leading to a conspicuous, roundish white blotch
in the centre of the leaf. The blotch is upper surface with full
depth spots - comprising of large clumps of frass.
. Puparium reddish |
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On Medicago, Melilotus and Trifolium in Britain and
in addition other genera of Fabaceae elsewhere. Common and widely
distributed in England. Also recorded in the Republic of Ireland.
Widespread and common throughout Europe.
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Agromyza
nana Meigen, 1830 [Diptera: Agromyzidae]. |
1e > Leaf-miner: Mine linear, whitish, both upper and lower surface. Pupation internal,
at the end of the mine with the anterior spiracles projecting through
the epidermis (Spencer, 1976:
433).
Upper-surface,
less often lower-surface corridor. Frass in isolated grains. Pupation
within the mine, usually in a lower-surface puparial chamber.
A long whitish upper surface corridor, which eventually goes lower surface. |
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Two
highly polyphagous species of Chromatomyia, with indistinguishable
mines, have been recorded in Britain. These are syngenesiae (Hardy) and horticola (Goureau) which can only be distinguished by the male genitalia. Both species are widespread in Britain and elsewhere, although syngenesiae is almost entirely restricted to Asteraceae. Records on Asteraceae not based on examination of male genitalia are treated in this account as Chromatomyia 'atricornis'.
Chromatomyia horticola is recorded on 160 plant genera in 31 families, of which 55 plant
genera in 19 families, but not yet on Trifolium, in Britain. On Trifolium elsewhere.
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Chromatomyia
horticola (Goureau, 1851) [Diptera: Agromyzidae]. |
1f > Leaf-miner: An upper surface linear mine with frass in conspicuous greenish
strips, largely alternating at each side of the channel. Puparium
yellow |
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Liriomyza congesta puparium
Image: © Willem Ellis (Bladmineerders van Europa) |
Polyphagous. On numerous genera of Fabaceae, including Trifolium, in Britain
and elsewhere. Widespread in Britain. Also recorded in the Republic
of Ireland and widespread and common throughout most of Europe
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Liriomyza congesta (Becker, 1903) [Diptera: Agromyzidae]. |
1g > Leaf-miner: The
mine starts as a long, narrow, winding corridor running towards
the midrib, widening to a blotch. Usually upper-surface, but in
small leaves also full-depth parts may occur. The blotch has broad
lobes; in their ends most frass is accumulated in the form of green
patches or clouds. Sometimes several larvae share mine. Pupation
usually in the soil, less often in the leaf (and then generally
not in the mine itself but in a small separated mine, that may even
be made in the petiole). |

Mine of Scaptomyza graminum on Cerastium glomeratum
Image: © Jean-Yves Baugnée (Bladmineerders van Europa) |
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On
? Amaranthus, Cerastium, Lychnis, Myosoton, Nasturtium, Silene, Stellaria, Atriplex,
? Anthyllis, ? Lupinus,
? Medicago, ? Montia and ? Antirrhinum, but not yet on Trifolium, in Britain.
On Amaranthus, Lepidium, Moricandia, ? Rorippa, Agrostemma, Arenaria, Cerastium, Corrigiola, Cucubalus, Dianthus, Gypsophila, Lychnis, Moehringia, Myosoton, Polycarpon, Saponaria, Silene, Spergularia, Stellaria, Vaccaria, Viscaria, Atriplex, Beta, Chenopodium, Obione, Salicornia, Spinacia, Anthyllis, Lupinus, Medicago, Allium, Montia, Portulaca and Antirrhinum elsewhere. Widespread in Britain and continental Europe.
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Scaptomyza
graminum (Fallén, 1823) [Diptera: Drosophilidae]. |
1h > Leaf-miner: Corridor-blotch
mine, normally dorsal; usually whitish; in small leaves it lies
characteristically in the centre of the leaf often touching the
petiole; in larger leaves it lies to one side of the mid-rib. Frass
deposited in green clumps near the leaf margin. Pupation usually
external, sometimes in a separate pupation mine.
Oviposition
whitin the leaf, at the lower surface. The first part of the mine
is a long, sometimes very long, corridor, that mostly will be overrun
by the later developments. Generally this corridor is directed,
frequently guided by a thick vein, to the midrib. The next section
of the mine is large, irregular blotch overlying the basal section
of the midrib. Locally large chunks of midrib tissue are consumed.
From this central blotch excursions are made into the leaf blade:
generally upper-surface, less often lower-surface and locally full-depth.
In plants with narrow leaves, like Diplotaxis, the mine may
occupy the entire width of the leaf. Often several larvae together
in a mine. Frass blackish green, powdery, in clouds, sometimes along
the sides of the corridors, later more in the periphery of the mine
and in the end of extensions of the blotch, sometimes seemingly
absent. (Often the frass can only be seen after the mine has been
opened). Pupation generally in the ground, rarely within he leaf,
in a short mine without frass. Hendel (1928a) described the biology,
larva and puparium.
An initial narrow gallery then leads to a blotch on the midrib of the leaf.
Watch a video of a scaptomyzid fly larva on Arabidopsis on YouTube by mash92587. |
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Polyphagous. On numerous genera of Brassicaceae, Asteraceae, Papaveraceae, Resedaceae, Tropaeolaceae and Violaceae, including Alliaria, Barbarea, Brassica, Cheiranthus, Cochlearia, Coincyia, Conringia, Diplotaxis, ? Eruca, Erysimum, ? Hesperis. Lepidium, Matthiola, Raphanus, ? Rorippa and Sinapis, but not yet on Trifolium, in Britain. On additional genera of these families and Fabaceae elsewhere. Widespread, from Caithness in the north to Cornwall in the south of Britain. Also recorded in the Republic of Ireland, Europe, the East Palaearctic, Near East and Neartic Region.
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Scaptomyza flava (Fallén, 1823) [Diptera: Drosophilidae]. |
1i > Leaf-miner: A short, irregular, linear upper surface mine on any part of the
leaf. Also recorded from young pods (Bland, 1997a).
Long
corridor mine. As a rule the first part of the mine is lower-surface,
the later part upper-surface. Often the loops are so dense that
a secondary blotch is the result. Because upper- and lower-surface
corridor segments often cross, the mine obtains a strange array
of transparant patches. There is no association with the midrib.
Frass in strings and thread fragments. Pupation outside the mine;
exit slit in upper epidermis.
Mine not associated with the veins or midrib of the leaf (It is this character which enables distinction from another Agromyzid pest species - Liriomyza huidobriensis). The larvae may leave one leaf (if not large enough) and enter another leaf, via the petiole). It exits the leaf to pupate through a semi-circular slit in the upper surface of the leaf. |
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Polyphagous. On 119 plant genera in 31 plant families of which only 4 plant genera in 2 plant families, but not yet on Trifolium, in Britain. Local, probably introduced
to Britain. Widespread in continental Europe particularly in Botanical
Gardens and glasshouses. Also recorded in Egypt.
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Liriomyza bryoniae (Kaltenbach, 1858) [Diptera: Agromyzidae]. |
Key for the identification of the known mines of British
non-Diptera recorded on
Trifolium
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Note:
The larvae of mining Coleoptera, Hymenoptera and Lepidoptera may live in a corridor mine, a corridor-blotch mine, a blotch mine, a case, a rolled or folded leaf, a tentiform mine or sandwiched between two more or less circular leaf sections in later instars. Larva may pupate in a silk cocoon. The larva may have six legs (although they may be reduced or absent), a head capsule and chewing mouthparts with opposable mandibles (see video of a gracillarid larva feeding). Larvae of Hymenoptera and Lepidoptera usually also have abdominal legs (see examples). Frass, if present, never in two rows. Unless feeding externally from within a case the larva usually vacates the mine by chewing an exit hole. Pupa with visible head appendages, wings and legs which lie in sheaths (see examples).
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1a > Leaf-miner and leaf-spinner: The larvae feed between spun leaves and often mine them. The mine is illustrated in British leafminers.
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On Genista, but not yet on Trifolium, in Britain. On Genista, Medicago and Trifoloium elsewhere. West and East Sussex and I. of Wight.
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Syncopacma vinella (Bankes, 1898) [Lepidoptera: Gelechiidae].
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1b > Leaf-miner: Small full depth blotch; older larvae free among spun leaves (Robbins, 1991a).
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On Lathyrus, Vicia and Trifolium in Britain and on Lathyrus, Melilotus and Trifolium elsewhere. Widespread in Britain. Also
recorded in the Republic of Ireland. Widespread in continental
Europe.
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Ancylis badiana (Denis & Schiffermmüller, 1775) [Lepidoptera: Tortricidae].
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1c > Leaf-miner: The
first generation initially forms an unmistakable leaf-mine on Anthyllis
vulneraria, but the second generation feeds on the flowers.
Feeding signs on other plants vary in appearance. Larvae can move
between sewn leaves, and more than one larva may be found together. Larvae
in a small full depth blotch, often with extensions. Frass concentrated
in one corner of the mine. The mining activities may cause the leaf
to roll inwards. Older larvae live free among spun leaves, but still
they may make then full depth mines by feeding on the leaf tissue
from a small opening. |
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Aproaerema anthyllidella larva, dorsal
Image: © Steve Wullaert (Bladmineerders van Europa) |
On Anthyllis, Medicago, Onobrychis, Ononis and Trifolium in Britain and Anthyllis, Chamaecytisus, Coronilla, Cysisus,
Dorycnium, Galega, Glycine, Hymenocarpos, Lathyrus,
Lotus, Medicago, Melilotus, Onobrychis, Ononis, Ornithopus, Oxytropis, Phaseolus, Psoralea, Trifolium, Trigonella and Vicia elsewhere. Britain including the Channel Is. and Northern Ireland.
Also recorded in the Republic of Ireland. Widespread in continental
Europe.
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Aproaerema
anthyllidella (Hübner, 1813) [Lepidoptera: Gelechiidae]. |
1d > Leaf-miner: In the first instar the larva mines the leaves, forming short,
irregular, blotch-like mines, but in later instars it lives externally,
feeding in spun leaves and often twisting those of tender shoots. Larval head light-brown or yellowish brown, edged with black postero-laterally,
ocellar area blackish; prothoracic plate black edged with whitish
anteriorly; abdomen dull dark green; pinacula distinct, black,
sometimes brownish but with black bases to setae; anal plate large,
black (Bradley et al., 1973).
Small,
full depth mine without a definite shape; little frass. Some silk
is deposited in the mine. The larva soon leaves the mine and continues
feeding among spun leaves. |
Polyphagous. On numerous genera and species of several plant families, but
not yet on Trifolium, in Britain. On numerous
genera and species of several plant families, including Trifolium,
elsewhere. Widespread in Britain and continental Europe. Also
recorded from the Channel Is.
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Cnephasia
incertana (Treitschke, 1835) [Lepidoptera: Tortricidae]. |
1e > Leaf-miner: The mine begins as a tiny, lower-surface epidermal corridor. Only
after the first moult the larva starts feeding on the leaf parenchyma.
The first result is a corridor overlying the midrib. From here branches
enter the leaf disk; gradually they widen and merge, laeving in
the end almost the entire leaf mined out. Almost all frass is ejected.
The larva can leave its mine and make a new one elsewhere. Pupation
witin the mine. |
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On Ononis and Trifolium in Britain and elsewhere.
Britain including East Kent and North Hants. Widespread in Continental
Europe.
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Parectopa
ononidis (Zeller, 1839) [Lepidoptera:
Gracillariidae]. |
1f > Leaf-miner: Lower surface tentiform mine that occupies only part of a leaflet.
In full grown leaves the mine is strongly inflated and is largely
hidden by the leaf. Unlike other species the entire leaflet is not
mined out.On Ononis and Trifolium and Lathyrus, Medicago,
Ononis, Trifolium and Vicia elsewhere. North Ebudes.
Also recorded in the Republic of Ireland. Widespread in continental
Europe.
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Phyllonorycter
insignitella (Zeller, 1846) [Lepidoptera: Incurvariidae]. |
1g > Leaf-miner: The mine is underside, occupying the whole leaflet, which turns
down at edges.
Lower
surface tentiform mine that occupies an entire leaflet; lower epidermis
strongly folded. Fully developed mines are strongly inflated and
the leaflet is so completely folded over the mine that the latter
becomes practically invisible. In this stage the larva has consumed
also all tissue in the roof of the mine, making the mine very conspicuous. |
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On Lathyrus, Medicago, Trifolium and Vicia in Britain
and Lathyrus, Lotus, Medicago, Trifolium and Vicia elsewhere. England, Ireland and continental Europe.
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Phyllonorycter
nigrescentella (Logan, 1851) [Lepidoptera: Gracillariidae]. |
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