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(Coleoptera, Diptera, Hymenoptera and Lepidoptera)
by
Brian Pitkin, Willem Ellis, Colin Plant and Rob Edmunds
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CARPINUS.
Hornbeam. [Betulaceae]
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Only
one species of Carpinus is recorded in Britain - Hornbeam
(C. betulus).
Twenty-four British miners are recorded on Carpinus.
A key to the European mines recorded on Carpinus
is provided in Bladmineerders van Europa.
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Hornbeam
Carpinus
betulus |
Key for the identification of the known mines of British
Diptera recorded on Carpinus
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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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1 > Leaf-miner: Larva
forming a narrow, unusually long, upper surface leaf-mine up to
12 cms, sometimes considerably widening at end; young leaves are
frequently distorted (Spencer,
1976: 93, fig. 130).
An
unusually long, upper-surface corridor that widens only little and
winds freely through the leaf. Frass in two neat rows. Pupation
outside the mine; exit slit (always?) in the lower epidermis.
A long, greenish, winding, upper surface gallery which sometimes broadens considerably at the end. |
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On Betula and Carpinus in Britain and only Betula elsewhere. Widespread in Britain. Also recorded in the Republic
of Ireland and continental Europe.
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Agromyza
alnibetulae Hendel, 1931 [Diptera: Agromyzidae]. |
Key for the identification of the known mines of British
non-Diptera recorded on Carpinus
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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 case-bearer: The larva lives outside the mine, protected by a case, and feeds on the underlying plant tissues via a hole cut in the epidermis. From that point it eats away as much leaf tissue as it can reach without fully entering the mine. Mine does not contain frass (Coleophora species) |
2 |
1b > Leaf-miner, but not a case-bearer: The larva lives mainly inside the mine. Mine usually contains frass. In later instars the larva may live sandwiched between two more or less circular sections cut from the leaf. |
3 |
2a > Leaf-miner
and case-bearer: The early case is tiny and the larva makes
a series of tiny holes on the leaf. After overwintering it makes
a shiny pistol shaped case in spring and window feeds British
leafminers).
The
young larva, before hibernation, makes tiny mines, sometimes tens
in one leaf. After hibernation window feeding is done. In this latter
stage the larva lives in a shining black pistol case of about 7
mm, that, with a mouth angle of 70°-80°, stands almost perpendicular
on the leaf. |
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On Crataegus, Malus, Prunus, Pyrus and Sorbus but not yet on Carpinus, in Britain plus numerous
genera and species of several plant families, including Carpinus,
elsewhere. Occurs in England and Wales, commoner in the south.
Widespread in continental Europe.
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Coleophora
anatipenella (Hübner, 1796) [Lepidoptera: Coleophoridae]. |
2b > Leaf-miner
and case-bearer: Larva mines leaves. The case is enlarged several times by mining
a leaf-edge and inserting the existing case into the mine which
is joined with silk. The final case is 7-8 mm long.
Larva
in a composite leaf case, composed of large leaf fragments. Characteristically,
the leaf fragments are attached in a failry untidy way. In spring
the case has two colours, because the old material (dull yellowish,
grey or pink) dates from before the hibernation, while new, reddish
brown material dates from after the winter. The case finally is
about 7-8 mm long; the mouth angle is 40-45°.. |
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On Alnus, Betula, Carpinus, Castanea and Corylus in Britain
and elsewhere. Widespread
in Britain and continental Europe, Also recorded in the Republic
of Ireland.
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Coleophora
binderella (Kollar, 1832) [Lepidoptera: Coleophoridae]. |
2c > Leaf-miner
and case-bearer: The full grown larva lives in a dull black pistol case of c 9 mm
that, with a mouth angle of 80-90° stands erect on the leaf.
Characteristic is the presence of some ear-like flaps. At least
after the hibernation the larvae do not mine any more, but rather
cause skeleton feeding. |

Case
of Coleophora currucipennella on Quercus rubra
Image: © Paul van Wielink (Bladmineerders van Europa) |
On Carpinus, Corylus, Quercus and Salix in Britain
and Betula, Carpinus, Corylus, Fagus, Quercus, Malus, Prunus,
Pyrus, Sorbus and Salix elsewhere. Widespread though
not common in Britain. Widespread in continental Europe.
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Coleophora
currucipennella Zeller, 1839 [Lepidoptera: Coleophoridae]. |
2d > Leaf-miner
and case-bearer: Lobe case. Many small leaf fragments are attached to the tubular
case; its end is strongly curved downwards. The larvae are attached
to the leaf underside, where they make a large number of relatively
small full depth mines. |
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On Betula and Corylus but not yet on Carpinus,
in Britain plus Alnus and Carpinus elsewhere. Widespread
but not common in Britain. Widespread in continental Europe. Also
recorded in the Republic of Ireland.
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Coleophora
fuscocuprella Herrich-Schäffer, 1855 [Lepidoptera:
Coleophoridae]. |
2e > Leaf-miner
and case-bearer: Larva mining leaves, the blotches brownish. The final case is 8-11
mm long, slender, and fixed at 45° to leaf surface, with anal
end laterally compressed. The case has a serrated keel due to formation
from the edge of a leaf.
Spatulate
leaf case. Strikingly slender, bivalved case, 8-11 mm long, with
a slight curve at the rear end, that is keeled and often toothed.
The end is laterally compressed. Mouth angle 45°. The full depth
mines often are conspicuously brown. |
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On Betula and Carpinus in Britain plus Alnus, Corylus and Myrica elsewhere. Widespread in Britain
and continental Europe. Also recorded in the Republic of Ireland.
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Coleophora
milvipennis Zeller, 1839 [Lepidoptera: Coleophoridae]. |
2f > Leaf-miner
and case-bearer: Composite leaf case. The material used to enlarge the case consists
of large pieces of full depth mine, that are attached with such
precision that they may seem seemless. In the course of summer an
autumn two (sometimes three, according to Hering, 1927b) pieces
are added. No more material is added after hibernation, causing
the case in spring to be rather uniformly coloured (contrary to C. binderella, that does add an extension in spring, and
is made of fresh leaf material). |

Case
of Coleophora orbitella on Betula pendula
Image: © Willem Ellis (Bladmineerders van Europa) |
On Alnus, Betula, Carpinus and Corylus in
Britain and elsewhere. Widespread in Britain and continental Europe.
Also recorded in the Republic of Ireland.
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Coleophora
orbitella Zeller, 1849 [Lepidoptera: Coleophoridae].
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2g > Leaf-miner
and case-bearer: The larva feeds by inserting its head into small mines it creates
on the leaves of birch, elm, alder, or hazel. Occasionally it is
found feeding on other trees, or on herbaceous plants onto which
it has accidentally Fallén. It forms two cases during its larval
life. The first case is initially curved, smooth, laterally compressed
with a bivalved anal opening, and about 2 mm long in September.
During October it feeds, and adds a few rough collars of larval
material around the oral opening. After hibernation, it feeds again
in April and early May, adding more protruding collars until they
equal or exceed the original smooth part of the case. At the same
time, it expands the case girth by the creation of a silk gusset
ventrally. The second case, 6 or 7 mm long, is formed in May, leaving
the vacated first case attached to its last feeding mine. The new
case is tubular with a trivalved crimp at the anal opening. The
dorsum is formed from the edge of the leaf from which the case was
cut. This results in a more or less serrated dorsal keel, depending
on the plant species and the individual piece of leaf used. Considerable
variation in the degree of serration can be found, even among specimens
off the same tree. The case colour varies with food plant, from
yellowish brown on birch, darkening through elm and hazel to dark
brown on alder.
The
strongly curved young case is is a composite leaf case, the adult
case is a tubular leaf case. The adult case is bivalved, about 7
mm in length; the mouth angle is around 30°. The case is straw
coloured and almost always has a toothed dorsal keel (remnant of
the margin of the leaf from which the case was cut). Neither larvae
or cases of C.
coracipennella, prunifoliae, serratella and spinella can be
separated; from serratella. |
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On Alnus, Betula, Corylus, Ulmus and Sorbus, but not yet on Carpinus, in Britain plus Carpinus, Mespilus,
Ostrya, Hippophae, Ribes, Myrica, Forsythia, Amelanchier, Chaenomeles,
Cotoneaster, Crataegus, Cydonia, Eriobotrya, Malus, Prunus, Sorbus,
Spiraea, Populus and Salix elsewhere. This is probably the commonest species of British
coleophorid, and is found throughout the British Isles. Widespread
in continental Europe.
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Coleophora
serratella (Linnaeus, 1761) [Lepidoptera: Coleophoridae]. |
2h > Leaf-miner
and case-bearer: Tubular leaf case. The case is almost barrel-shaped, with a large
leaf fragment that, while withering, folds itself untidily arround
the tube.
A
biennial life cycle in the UK (may be annual in continental Europe). The second
and third cases are formed by cutting out a large leaf portion and
then wrapping it around - leaving an edge protruding, which then
withers. |
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On Betula, Crataegus, Malus and Sorbus but not yet on Carpinus, in Britain plus Alnus, Carpinus, Malus,
Sorbus and Tilia elsewhere. Widespread but not common
in Britain. Also recorded in the Republic of Ireland. Widespread
in continental Europe.
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Coleophora
siccifolia Stainton, 1856 [Lepidoptera: Coleophoridae]. |
2i > Leaf-miner
and case-bearer: The larva feeds on a wide range of trees, shrubs and herbs, favouring
Rosaceae, but not exclusively. The fully developed cased larva may
be found active in October and again, after winter diapause, in
April. Cases, about 6 mm, of diapausing larvae may be found through
winter, fixed to a tree or fence post. The dorsal surface of the
case is usually covered in leaf fragments, but they can sometimes
be worn off almost smooth. The ventral surface is swollen at the
middle and has a keel, which usually bends upwards at the posterior.
The cases of C. ahenella (on Rhamnus, Frangula, Viburnum and Cornus) and C.
potentillae (case less swollen, keel not bent up, resting
position less prone) are very similar.
Brownish
lobe case that lies almost flat on the leaf, either on the upper
or on the lower side. Case widest about the middle. Ventrally there
is a distinct keel. Mouth angle 0°. Full depth mines rather
large. The flaps of cuticular tissue that serve to enlarge the case
are cut out of the upper epidermis. (contrary to C.
ahenella and C.
potentillae, that use tissue from the lower epidermis).
The removal of these tissue flaps creates holes that are much larger
than those that serve as the entrance to the mine. |
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Coleophora violacea larva, lateral
Image: © Willem Ellis (Bladmineerders van Europa) |
Polyphagous. On numerous genera and species in several plant families, including Carpinus, in Britain and elsewhere. Widespread in Britain
and continental Europe.
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Coleophora
violacea (Ström, 1783) [Lepidoptera: Coleophoridae]. |
3a > Leaf-miner: The larvae mine the leaves at first, then create small feeding windows.
Small,
full depth, hook-like corridor, usually in a vein axil, with a proportionally
large larval chamber. The remainder of the mine almost entirely
stuffed with frass. At the start if the mine an iridescent egg shell.
The larvae soon leave their mine and start living free on the leaf. The pupa and white ribbed cocoon are illustrated in British
leafminers. |
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Bucculatrix thoracella cocoon
Image: Rob Edmunds (British leafminers) |
On Tilia, but not yet on Carpinus, in Britain and Acer, Aesculus, Betula, Carpinus, Fagus, Sorbus and Tilia elsewhere. Widely distributed in southern England. Widespread
in continental Europe.
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Bucculatrix
thoracella (Thunberg, 1794) [Lepidoptera: Bucculatricidae].
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3b > Leaf-miner: The larvae mine leaves at first, forming a blotch mine, later descending
to the ground in a portable case and feeding on dead leaves.
Oviposition
is by way of an ovipositor, therefore no egg shell visible. The
larva makes a small, roundish, blotch; often several in a leaf.
Already after its first moult it makes an excision out of the mine,
in size almost equal to the blotch (3-4 mm). Thus sandwiched it
drops to the ground and continues feeding on dead leaf material. |

Mines
of Incurvaria masculella
Image: © Rob Edmunds (British
leafminers) |
On Crataegus and Rosa but not yet on Carpinus,
in Britain and Carpinus, Corylus, Vaccinium, Catanea, Fagus,
Quercus, Crataegus, Rosa and Tilia elsewhere. Widespread
in Britain, Ireland and continental Europe.
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Incurvaria
masculella (Denis & Schiffermüller, 1775) [Lepidoptera:
Incurvariidae]. |
3c > Leaf-miner: Oviposition by way of an ovipositor, no egg visible therefore.
The larva makes an irregular blotch. The part of the mine nearest
to the oviposition site is more thranslucent than the later, in
transparancy more greenish, part of the mine. The mine usually lies
close to the leaf tip, often several together. After its first moult
the larva makes a roundish excision, 3-4 mm in diameter. Incurvaria larvae, while resting, take a horse-shoe like posture, unlike the larvae of Antispila species. Sandwiched herein it drops to the ground and continues feeding of dead leaves.
The excision occupies about half of the surface of the blotch.
The
mine is also described in. |
On Vaccinium myrtillus, Prunus and Rubus chamaemorus, but not yet on Carpinus, in Britain. On
several genera and species of several plant families, including Carpinus, elsewhere. Widespread in much of the British Isles
and continental Europe.
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Incurvaria
oehlmanniella (Hübner, 1796) [Lepidoptera: Incurvariidae]. |
3d > Leaf-miner: The larva starts making a corridor of a few mm, followed, and
mostly overrun, by a circular blotch of 4-5 mm diameter.
Generally
several larvae feed in a single leaf, creating a distinctive pattern
of feeding windows. The larvae then cut out circular cases and drop
to the leaf-litter to continue feeding, leaving behind a leaf containing
many circular or oval cut-outs. |
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On Alnus, Betula, Carpinus, Corylus, Malus and Tilia, but not yet on Carpinus, in Britain. On
several genera and species in several plant families, including Carpinus, elsewhere. Fairly well-distributed throughout
much of the British Isles, though it tends to be commoner further
north. Widespread in continental Europe.
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Incurvaria
pectinea Haworth 1828 [Lepidoptera: Incurvariidae]. |
3e > Leaf-miner: The
larva feeds on hazel or hornbeam, creating blotches with intertwining
threads of frass, typical of the genus.
Large
white blotch, starting at the leaf margin. Frass in long threads.
Often several larvae in a mine. Pupation outside the mine. |
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On Carpinus and Corylus in Britain plus Alnus and Ostrya elsewhere. Widespread in England and recorded
in the Republic of Ireland. Widespread in continental Europe.
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Eriocrania chrysolepidella (Zeller, 1851) [Lepidoptera: Eriocraniidae]. |
3f > Leaf-miner: The
initial mine expands to form a full depth blotch. It resembles Phyllonorycter
tenerella, but has a mottled lower surface. It then forms
two folds.
Small,
angular, full depth blotch, often in a vein axil. Lower, in the
end also upper, epidermis brown. The larva deposits some silk in
the mine, but the quantity is so low that the mine remains practicaly
flat. Later the larva leaves the mine and continues feeding within
a downfolded leaf margin or leaf tip. |
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On Carpinus betulus in Britain. On Acer platanoides,
Acer pseudoplatanus, Carpinus betulus, Carpinus orientalis and Ostrya carpinifolia elsewhere. South-east England.
Widespread in continental Europe.
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Parornix
carpinella (Frey, 1863) [Lepidoptera: Gracillariidae]. |
3g > Leaf-miner: The mine is upper side, over veins. Silvery, with brown speckling,
later contracting to cause leaf to fold upwards.
Upper-surface
tentiform mine. The early mine is roundish, silvery, flat, and lies
centered over a side vein. The older mine strongly contracts and
sometimes almost doubles the leaf. |
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On Carpinus betulus in Britain and Carpinus betulus and Ostrya carpinifolia elsewhere. A local species, mainly
found in the south and south-east of England northwards to the
midlands and South York, the most northerly record to date. Widespread
in continental Europe.
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Phyllonorycter
esperella (Goeze, 1783) [Lepidoptera: Gracillariidae]. |
3h > Leaf-miner: The mine is oval on Q. ilex (note - there may be several
mines in the leaf), and similar to P.
quercifoliella on deciduous oaks. It is between adjacent
veins on beech and hornbeam.
Small,
oval, lower-surface tentiform mine, 9-14 mm long, mostly between
two lateral veins. The lower epidermis with a single sharp fold
(sometimes forked near its end). Pupa in very flimsy cocoon, that
contains a bit of frass laterally and at the rear end. |
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On Betula, Carpinus, Castanea, Fagus, Nothofagus, Quercus,
Malus, Ostrya and Prunus in Britain and Carpinus, Castanea,
Fagus, Quercus, Prunus and Tilia elsewhere. Widespread
in Britain, Ireland and continental Europe.
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Phyllonorycter
messaniella (Zeller, 1846) [Lepidoptera: Gracillariidae].
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3i > Leaf-miner: The mine is oval, sometimes crossing veins. The lower epidermis
with several strong creases.
Along
with P. tristrigella,
this species feeds on elm, although not usually on wych elm like
that species. However the two mines are quite different - schreberella forming a rounded mine and tristrigella a longer, narrower tube-like
mine.
Short,
roundish to oval, somewhat inflated, lower surface tentiform mine,
often crossing a lateral vein. Epidermis with several length folds. Dark brown pupa in a tough, greenish cocoon; the cocoon lies
free in the mine, and its wall is not encrusted with frass. |
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On Ulmus but not yet on Carpinus, in Britain
and ? Carpinus and Ulmus elsewhere. Mainly in the
southern half of England and Wales. Widespread in continental
Europe.
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? Phyllonorycter schreberella (Fabricius,
1781) [Lepidoptera: Gracillariidae]. |
2j > Leaf-miner: A mine between veins from midrib to leaf-edge, narrow, tubular,
with one crease in lower epidermis.
Narrow,
finally tubular lower-surface mine between two side veins. The lower
epidermis with one strong fold. Pupa in a flimsy cocoon in a corner
of the mine, usually in the axil of midrib and side vein. Frass
loosely heaped in the opposite corner. |
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On Carpinus in Britain and Carpinus and Ostrya elsewhere. Southern half of England. Widespread in continental
Europe.
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Phyllonorycter
tenerella (Joannis, 1915) [Lepidoptera: Gracillariidae]. |
3k > Leaf-miner: Oviposition at the underside of the leaf, in the axil of a vein.
The mine is a not very slender corridor. The first part is almost
filled with frass; in the later part the frass lies in thick lumps.
The trajectory of the mine is not angular, neither is it determined
by the leaf venation. The discrimination between this mine and the
one of Stigmella
floslactella is difficult. |

Mine
of Stigmella carpinella on Carpinus betulus
Image: © Willem Ellis (Bladmineerders van Europa) |
On Carpinus in Britain and Carpinus and Ostrya elsewhere. West Kent in Britain. Widespread in continental Europe.
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Stigmella
carpinella (Heinemann, 1862) [Lepidoptera: Nepticulidae]. |
3l > Leaf-miner: The early gallery is filled with frass, later leaving clear
margins.
Like Stigmella microtheriella the larva feeds on hazel or hornbeam, and its mines are often found
alongside that species in the same leaf. However the mines of S.
floslactella are generally wider, less angular and contain more
scattered frass than those of S. microtheriella.
Oviposition
at the underside of the leaf, in a vein axil. Mine a slender, gradually
widening corridor; the last section is clearly wider than the larva.
In the first section the vaguely delimited frass line almost fills
the corridor. Later the frass lies in irregular arcs and clouds,
filling about one third of the width of the corridor. The trajectory
of the mine is not angular, independent of the leaf venation. Pupation
external, exit slit in the upper epidermis. |
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On Betula, Carpinus and Corylus in Britain and Carpinus,
Corylus and Ostrya elsewhere. Widespread in Britain
and continental Europe. Also recorded in the Republic of Ireland.
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Stigmella
floslactella (Haworth, 1828) [Lepidoptera: Nepticulidae]. |
3m > Leaf-miner: A narrow gallery, tending to follow veins of leaf. The early part
with linear frass.
Oviposition
at the underside of the leaf, mostly close to a vein. The mine is
a long, very slender corridor; even towards the end hardly wider
than necessary to accomodate the growing larva. Frass in a narrow
central line. The shape of the mine differs somewhat between the
hostplants. In Carpinus the mine closely follows a heavy
vein over a long distance; also the mine tends to be somewhat shorter
and broader, and the frass often lies in a more diffuse line. The
mines in Corylus are not so strictly defined by the venation
and the frass line is narrower (Emmet, 1983a; Johansson ao, 1990a).
Sometimes it is difficult to separate the mines from those of S.
floslactella; an additional difference then is that even
in the very first part of the corridor the frass of microtheriella lies in a narrow line, while the frass of floslactella seems
to fill the entire corridor there. The pale golden larva lies venter-upwards
in the mine. Sometimes there can be several larvae mining the
same leaf. |
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On Carpinus betulus, Corylus avellana, Nothofagus and Ostrya carpinifolia in Britain and Carpinus spp.
and Ostrya spp. elsewhere. Widespread in Britain, Ireland
and continental Europe.
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Stigmella
microtheriella (Stainton, 1854) [Lepidoptera: Nepticulidae]. |
3n > Leaf-miner: Full depth blotch, invariably beginning at the leaf tip or the tip
of a leaf lobe or tooth. Oviposition site covered by a black, shining
drop of hardened secretion. Frass generally in long threads, but
sometimes in elongated granules. Pupation in the mine, not in a cocoon. |
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On Corylus avellana, but not yet on Carpinus,
in Britain. On several genera and species in several
plant families including Carpinus elsewhere. Widespread
in Britain and continental Europe.
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Trachys
minutus (Linnaeus, 1758) [Coleoptera: Buprestidae]. |
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