Subject Editor: Maria Heikkilä
The leafmining family
In Europe,
While a predecessor of the present manuscript was in preparation, a slightly worn small female moth with a six mm wingspan and a golden median fascia with two other golden forewing spots was caught, on 23.viii.2016, in an MV Robinson moth trap in the Wildlife Garden of the Natural History Museum in South Kensington (UK, London). DCL identified this moth provisionally as an
Here we analyse material from throughout Europe and show that
Details of specimens examined are provided in the Suppl. material
Measurements of genitalia and forewing details (Table
Material of
Identification |
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Sample ID | Process ID | Hostplant | GenBank | Institution |
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HELA015–13 |
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HELA016–13 |
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BVS03 | ANTVI029–11 |
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ANTVI009–11 |
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HELA023–13 |
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HELA024–13 |
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TER1CS | ANTVI035–11 |
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TER2CS | ANTVI036–11 |
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HELA025–13 |
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HELA142–16 |
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HELA143–16 |
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DL-AP1 | HELA150–17 |
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CLV2597 | GRPAL724–11 |
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INRA |
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TER3CM | ANTVI037–11 |
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ANTVI040–11 |
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ANTVI038–11 |
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ANTVI039–11 |
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ANTVI019–11 |
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HELA029–13 |
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LEAFN053–13 |
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LEAFN054–13 |
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LEAFN055–13 |
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LEAFN079–13 |
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LEAFN389–13 |
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LEAFN390–13 |
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LEAFN391–13 |
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HELA144–16 |
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HELA145–16 |
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DL-AT1 | HELA148–17 |
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DL-AT2 | HELA149–17 |
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DL-AT3 | HELA147–17 |
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HELA034–13 |
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HELA033–13 |
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HELA047–13 |
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COPIN003–14 |
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MM15506 | LEFIG642–10 |
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MM15507 | LEFIG643–10 |
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MM06188 | LEFID308–10 |
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HELA058–13 |
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MM05954 | LEFIA1161–10 |
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MM05955 | LEFID174–10 |
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HELA102–14 |
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AVS09 | ANTVI027–11 |
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HELA107–14 |
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The sequence data generated and used in this study have been deposited in the public BOLD dataset (“
Comparison of measurements of forewing and costal spot in
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Range | Mean ± sd | # | Range | Mean ± sd | # | |
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Forewing length | 2.19–2.70 mm | 2.52 ± 0.19 | 7 | 2.54–3.11 mm | 2.78 ± 0.19 | 11 |
Costal spot width | 0.23–0.39 mm | 0.33 ± 0.07 | 7 | 0.35–0.44 mm | 0.37 ± 0.03 | 11 |
Costal spot length | 0.34–0.56 mm | 0.44 ± 0.07 | 7 | 0.36–0.45 mm | 0.41 ± 0.03 | 11 |
Ratio spot l/w |
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7 |
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Ratio spot w/fw l | 0.098–0.148 | 0.129 ± 0.019 | 7 | 0.116–0.157 | 0.135 ± 0.014 | 11 |
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Forewing length | 2.16–2.91 mm | 2.61 ± 0.23 | 7 | 2.52–2.82 mm | 2.65 ± 0.10 | 10 |
Costal spot width | 0.25–0.44 mm | 0.35 ± 0.06 | 7 | 0.35–0.44 mm | 0.40 ± 0.03 | 10 |
Costal spot length | 0.35–0.46 mm | 0.42 ± 0.04 | 7 | 0.32–0.43 mm | 0.39 ± 0.03 | 10 |
Ratio spot l/w |
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Ratio spot w/fw l | 0.110–0.167 | 0.132 ± 0.019 | 7 | 0.124–0.169 | 0.151 ± 0.013 | 10 |
In Europe, there are three widespread native species of
For
In our experience, many records of hostplants of
Taxa | Locality | Host-plant | Original identification | Reference |
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UK |
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UK, Surrey, Priest Hill, |
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DCL reared | |
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HU: Pécs, 17.xi.1971, 27.x.1974 |
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HU: Budapest, Budakeszi, leg. Szőcs | [interpreted as |
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PL: Woj. Poznan | Not given |
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CZ: Koda near Beroun, 3.1954 |
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CZ: Koda near Beroun, 3.1954 |
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“a leafminer” [interpreted as |
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CZ |
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HU: Normafa, 14.vii.1954 | [interpreted as |
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UK, SE London, Kelsey Park, x.2016 |
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DCL reared | |
UK: London, Victoria Embankment, 30.vi.2017 |
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DCL reared | |
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NL: Simpelveld [as Simpelweld] | Not given |
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PL: Torún |
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UK |
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Not given |
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HU: Pécs, Üszög,1.x.1973 |
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DE: München | Not given |
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[primary source not seen] | Not given |
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[primary source not seen] | Not given |
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In the female genitalia,
1.
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3.
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5.
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7.
Provisionally in
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* species introduced from North America
1 | Forewing shiny grey to brown, with only one or two spots along dorsum | |
– | Forewing with pale spots at both costal and dorsal margin, often a fascia present as well |
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2 | Forewing basally silvery white, distally dorsal area black and costal area yellow, with two conspicuous triangular silvery spots. Very small, wingspan 3.5–4.7 mm, introduced on |
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– | Forewing more or less uniform brown or fuscous, with a pale fascia, sometimes broken, at 1/3 and two opposite spots at 2/3 |
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3 | Forewing with an additional white silvery spot in apex, not touching the wing margins. Head silvery white. Wingspan 5.5–6.2 mm. Introduced on |
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– | Forewing without an additional apical spot, head not white, various tinges of grey, brown or leaden. Native species |
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4 | Small species, wingspan 4.0–4.7 mm. Venation reduced, no cell in forewing. Antenna with 15–20 segments. Leafminers on |
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– | Larger species, wingspan at least 5.0 mm, usually more. Venation complete with discoidal cell (Figs |
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5 | Forewing fascia usually divided into two spots, spots relatively large, antenna with 15 segments. Male foretibia with small epiphysis. Mediterranean species, mines on |
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– | Forewing fascia not divided in two spots, spots small, antenna with 20 segments. Male foretibia without epiphysis. Rare Central European species, mines on |
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6 | Smaller species, wingspan 4.8–7.0 mm; forewing and thorax dark fuscous to black, head greyish; male forewing underside with bunch of yellow androconial scales (Figs |
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– | Larger species, wingspan 6.8–8.5 mm, usually larger than 7.0 mm; wings and thorax coppery, head golden; male forewing underside without androconial scales. Antenna with ca 25 segments. |
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7 | Forewing costal spot usually triangular, ratio length/width: 1.18–1.63 in males, 1.05–1.52 in females; slightly smaller, wingspan 4.8–6.1 mm. Best distinguished by male genitalia, larva and hostplant: |
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– | Forewing costal spot usually trapezoid to almost square, ratio length/width: 0.93–1.22 in males, 0.78–1.17 in females; slightly larger, wingspan 5.7–7.0 mm. Best distinguished by male genitalia, larva and hostplant: |
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1 | Uncus medially blunt, central lobe reduced or absent. Valva at inner margin with spinelike process (Figs |
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– | Uncus with prominent central lobe. Valva without spinelike process |
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2 | Horseshoe-shaped anellus present between the valvae (Fig. |
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– | No anellus between valvae visible (Fig. |
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1 | Near start of mine 2–7 small brown scars made by ovipositing female (“test punctures”) (Figs |
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– | No scars near start of mine. Larva with row of dorsal black sclerotised plates (note: larvae usually feed with ventral side upwards) (Figs |
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2 | Mine on |
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– | Mine on |
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Total 23♂, 27♀: France (leafmines), Germany (2♂, 1♀, leafmines), Greece (1♂, larvae, leafmines), The Netherlands (14♂, 12♀, larvae, leafmines), Switzerland (5♂, 14♀, larvae, leafmines), United Kingdom (1♂, larvae). Details in Suppl. material
Larva easily separated from
Male (Figs
Female (Fig.
Measurements, male: forewing length 2.3–2.9 mm (2.7 ± 0.2, 12), wingspan 4.8–6.1 mm, 19–20 antennal segments (n=8); female: forewing length 2.3–3.0 mm (2.7 ± 0.2, 7), wingspan 4.8–6.1 mm, 19–20 antennal segments (n=3). For costal spot see Table
Male genitalia (Figs
Female genitalia (Figs
Larva (33–35, 39, 40). Pale grey translucent, head capsule brown, prothorax with large black tergum and sternum. In instar IV, the final feeding instar, mesothorax, metathorax and abdominal segments 1 to 7 dorsally each with a central black spot, with fuzzy outline, more or less rhomboid, spots becoming smaller from segment 5 to 7; ventrally up to 5 spots on metathorax and segments 1 to 4. Abdominal segment 8 dorsally with a swollen hump, at the anal end lined with a single row of 5 black warts. Anal segment black. More details on earlier instars are given by
Widespread in Europe, local in southern England, in the Netherlands and Belgium local in hilly limestone areas in the South and East. Throughout central and southern Europe, but not known in detail, due to confusion with
The natural distribution of
This species was described from an unspecified number of specimens reared by Martini from mines on
Martini’s study is very thorough, comparing
The male genitalia figured by Wojtusiak and reproduced by Razowski resemble more
Total 34♂, 33♀, 4 sex unknown: Albania (1 ♀), Austria (2♂, 2♀, 2 sex unknown), Bulgaria (larvae, mines), Czech Republic (6♂, 3♀, leafmines), France (leafmines), Germany (1♂, 1♀, leafmines), The Netherlands (16♂, 16♀, 2 sex unknown, larvae, leafmines), Poland (4♂, 2♀), Switzerland (5♂, 7♀, larvae, leafmines), United Kingdom (1♀, larvae, leafmines). Details in Suppl. material
See
Male (Fig.
Female (Fig.
Measurements, male: forewing length 2.7–3.3 mm (2.9 ± 0.2, 13), wingspan 5.7–7.0 mm, 19–20 antennal segments (n=8); female: forewing length 2.3–3.0 mm (2.7 ± 0.2, 7), wingspan 5.7–7.0 mm, 19–20 antennal segments (n=3). For costal spot see Table
Male genitalia (Figs
Female genitalia (Figs
Larva (Figs
There are some records on the planted species
Widespread in central and southern Europe, in the natural range of
The natural distribution of
The original description by Fischer von Röslerstamm (1843) of
Fischer von Röslerstamm’s Microlepidoptera were acquired in 1843 by Herrich-Schäffer (
It is somewhat mysterious why Dziurzyński needed to introduce
Total 8♂, 20♀: Austria (2♂, 2♀), Bulgaria (larva, leafmines), France (1♀, larva, leafmines), Germany (2♀), The Netherlands (4♂, 13♀, larvae, leafmines), Poland (1♂, 1♀), Romania (leafmines), Switzerland (1♂, 1♀). Details in Suppl. material
(male). forewing length 3.7–3.9 mm (3.8 ± 0.1, 5), wingspan 7.8–8.5 mm, 24–25 antennal segments; female: forewing length 3.3–4.0 mm (3.7 ± 0.2, 11), wingspan 6.9–8.7 mm, 24–25 antennal segments.
Widespread in central and southern Europe, distributed further north than the other species: occurs in a few localities in southern Norway and southern Sweden, the islands Gotland and Öland, the islands of Denmark, Estonia, Latvia, Lithuania, southern England to Midlands, all West and Central European Countries, just in the NE of Spain (
Hübner (1813) used the name
We included all ten European species of
Maximum likelihood tree of COI barcodes of 43 European
Identifying material based on DNA barcodes is highly reliable for this group and provides an alternative to morphology. True
About 20 species or cultivars of
At Kew and Wisley, living specimens labelled as
At Cambridge Botanical Garden in October 2016, all
As
In Wisley, the 14 specimens of
At
At Kew Gardens herbarium, only four specimens of
The herbarium of the Naturalis Biodiversity Center, Leiden was briefly scanned by means of its BioPortal (
Our results show that the
Literature and other records of
All records of either species from the reciprocal principal hostplant need to be looked at with suspicion: whereas we do not a priori exclude the possibility, probably the majority are either a case of misidentified hostplant or misidentified leafminer: we did not find any convincing case of host switching between them, and we could reliably re-identify hosts of several online records. The fact that the two European
In summary, all our results support the hypothesis that the two species
The first reports of a range expansion of
The next records are from an expansion in The Netherlands in 1996 and 1997 (Kuchlein and van Frankenhuyzen 1999), then considered to represent a sudden expansion of the native populations that live in the south of the province of Limburg on
In Germany the expansion was noted a bit later, and the species was recorded as new for Sachsen in Belgershain in 1999 by Stübner (
In the UK, the earliest confirmed specimen still remains the adult of
We have no indication of expansion of
In the Netherlands
While many species have been observed in recent years to spread over Europe, whether they are native in Europe or not, the expansion of
The expansion of
It is certainly possible that climate change also has played a role in the expansion of the leafmining moths, but we have no data supporting this.
With this paper in hand it should be easy to recognise the mines of the three
Many people helped us with information or material, we are grateful for their help: Willem Ellis collected additional
Specimen and Locality Data
Excel file
Explanation note: Specimen data.
Figure S1.
Figure S2.
Adobe PDF file.
Explanation note: Tree topology for the phylogenetic hypothesis adopted, to be used as input in applications reading nexus (requires some slight previous edition).