Research Article |
Corresponding author: Erik J. van Nieukerken ( nieukerken@naturalis.nl ) Academic editor: Maria Heikkilä
© 2018 Erik J. van Nieukerken, David C. Lees, Camiel Doorenweerd, Sjaak (J.C.) Koster, Rudolf Bryner, Arnold Schreurs, Martijn J.T.N. Timmermans, Klaus Sattler.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
van Nieukerken EJ, Lees DC, Doorenweerd C, Koster S(JC), Bryner R, Schreurs A, Timmermans MJTN, Sattler K (2018) Two European Cornus L. feeding leafmining moths, Antispila petryi Martini, 1899, sp. rev. and A. treitschkiella (Fischer von Röslerstamm, 1843) (Lepidoptera, Heliozelidae): an unjustified synonymy and overlooked range expansion. Nota Lepidopterologica 41(1): 39-86. https://doi.org/10.3897/nl.41.22264
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Antispila treitschkiella (Fischer von Röslerstamm, 1843) and A. petryi Martini, 1899, sp. rev. were regarded as synonymous since 1978, but are shown to be two clearly separated species with different hostplants, life histories, DNA barcodes and morphology. Antispila treitschkiella feeds on Cornus mas L., is bivoltine, and has, by following its ornamentally planted host, greatly expanded its range in north-western Europe. In contrast A. petryi feeds on the widespread native C. sanguinea L., is univoltine, and is one of only two Antispila species previously resident in the British Isles, the Netherlands and northern Europe. Consequently, the increase in abundance of A. treitschkiella in the Netherlands since the early 1990s and in Great Britain in recent years must be regarded as part of a recent expansion into north-western Europe, whereas the native A. petryi is hardly expanding and less abundant. In Britain, detailed surveys of parks and living collections confirmed the monophagy of these two species. A search of British herbarium samples provided no evidence for an earlier date of establishment. Information on recognition of all stages, including DNA barcodes, and distribution is provided, and these two species are compared with the third European Cornus L. leafminer, A. metallella (Denis & Schiffermüller, 1775).
The leafmining family Heliozelidae has about 125 named species and is especially diverse in Australia and the Americas (
In Europe, Heliozelidae are poorly represented, with only ten species recorded, including two recent invasives from North America: Antispila oinophylla van Nieukerken & Wagner, 2012 and Coptodisca lucifluella (Clemens, 1860) (van
Antispila treitschkiella was reported to expand suddenly in distribution and abundance on planted C. mas in parks in the Netherlands (
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 Antispila species, a genus not previously reported from the South Kensington area. This determination was confirmed by EvN from a photograph, a finding that struck DCL as peculiar since the Antispila species known in Britain generally fly in May–June (A. metallella) or June–July (A. treitschkiella sensu auctorum). The specimen was too small to correspond to A. metallella, nor had it the typical size and markings, thus ruling out this species. Surprisingly, no mines could be found on C. sanguinea in the NHM Wildlife Garden, nor on adjacent beds containing C. alba L. A few vacated mines were subsequently found on old plants of C. sanguinea in nearby Kensington Gardens. This find led later to the discovery of many C. mas feeding Antispila in London.
Here we analyse material from throughout Europe and show that A. treitschkiella as currently used is a composite species: A. petryi should be re-instated as a valid, well separated species. In order to recognise the three European Cornus feeding species, they are briefly redescribed and diagnostic differences are presented.
Material. Much of the material for this paper was collected or observed by the authors; additional specimens from the following collections are considered.
UPI University of Padova, Dept. of Environmental Agronomy and Crop Science, Italy
Details of specimens examined are provided in the Suppl. material
Rearing. Collected leaves were kept in polystyrene jars or bags, with some moss and/or paper tissue added, until the larvae had prepared their leaf-epidermis-shield cases in the fourth instar. It was often necessary to remove the cut-out cases manually from the leaves, after which the leaves were taken from the rearing jars and dried as vouchers.
Morphology. Methods for preparation of the genitalia follow
Measurements of genitalia and forewing details (Table
Material of Heliozelidae used for DNA barcodes, all in dataset DS-ANTITR, doi: 10.5883/DS-ANTITR.
Identification | BIN | Sample ID | Process ID | Hostplant | GenBank | Institution |
---|---|---|---|---|---|---|
Antispila metallella | ACG8679 | RMNH.INS.18463 | HELA015–13 | Cornus sericea | MG191951 | RMNH |
Antispila metallella | ACG8679 | RMNH.INS.24371 | HELA016–13 | Cornus sericea | KP697814 | RMNH |
Antispila oinophylla | AAI4367 | BVS03 | ANTVI029–11 | Vitis vinifera | JQ412531 | UPI |
Antispila oinophylla | AAI4367 | RMNH.INS.24038 | ANTVI009–11 | Vitis vinifera | JQ412537 | RMNH |
Antispila petryi | AAV5055 | RMNH.INS.18803 | HELA023–13 | Cornus sanguinea | MG191953 | RMNH |
Antispila petryi | AAV5055 | RMNH.INS.24372 | HELA024–13 | Cornus sanguinea | MG191950 | RMNH |
Antispila petryi | AAV5055 | TER1CS | ANTVI035–11 | Cornus sanguinea | JQ412517 | UPI |
Antispila petryi | AAV5055 | TER2CS | ANTVI036–11 | Cornus sanguinea | JQ412518 | UPI |
Antispila petryi | AAV5055 | RMNH.INS.29304 | HELA025–13 | Cornus sanguinea | MG191937 | RMNH |
Antispila petryi | AAV5055 | RMNH.INS.30614 | HELA142–16 | Cornus sanguinea | MG191936 | RMNH |
Antispila petryi | AAV5055 | RMNH.INS.30615 | HELA143–16 | Cornus sanguinea | MG191947 | RMNH |
Antispila petryi | AAV5055 | DL-AP1 | HELA150–17 | Cornus sanguinea | KY613518 | NHMUK |
Antispila treitschkiella | AAU1917 | CLV2597 | GRPAL724–11 | Cornus mas | MG521913 | INRA |
Antispila treitschkiella | AAU1917 | TER3CM | ANTVI037–11 | Cornus mas | JQ412553 | UPI |
Antispila treitschkiella | AAU1917 | RMNH.INS.11818 | ANTVI040–11 | Cornus mas | JQ412554 | RMNH |
Antispila treitschkiella | AAU1917 | RMNH.INS.11856 | ANTVI038–11 | Cornus mas | JQ412555 | RMNH |
Antispila treitschkiella | AAU1917 | RMNH.INS.11857 | ANTVI039–11 | Cornus mas | JQ412551 | RMNH |
Antispila treitschkiella | AAU1917 | RMNH.INS.11858 | ANTVI019–11 | Cornus mas | JQ412552 | RMNH |
Antispila treitschkiella | AAU1917 | RMNH.INS.29300 | HELA029–13 | Cornus mas | MG191955 | RMNH |
Antispila treitschkiella | AAU1917 | RMNH.INS.544573.1 | LEAFN053–13 | Cornus mas | MG191949 | RMNH |
Antispila treitschkiella | AAU1917 | RMNH.INS.544573.2 | LEAFN054–13 | Cornus mas | MG191954 | RMNH |
Antispila treitschkiella | AAU1917 | RMNH.INS.544573.3 | LEAFN055–13 | Cornus mas | MG191952 | RMNH |
Antispila treitschkiella | AAU1917 | RMNH.INS.544585 | LEAFN079–13 | Cornus mas | MG191941 | RMNH |
Antispila treitschkiella | AAU1917 | RMNH.INS.552214.1 | LEAFN389–13 | Cornus mas | MG191945 | RMNH |
Antispila treitschkiella | AAU1917 | RMNH.INS.552214.2 | LEAFN390–13 | Cornus mas | MG191939 | RMNH |
Antispila treitschkiella | AAU1917 | RMNH.INS.552214.3 | LEAFN391–13 | Cornus mas | MG191944 | RMNH |
Antispila treitschkiella | AAU1917 | RMNH.INS.30611 | HELA144–16 | Cornus mas | MG191942 | RMNH |
Antispila treitschkiella | AAU1917 | RMNH.INS.30613 | HELA145–16 | Cornus mas | MG191948 | RMNH |
Antispila treitschkiella | AAU1917 | DL-AT1 | HELA148–17 | Cornus mas | KY613515 | NHMUK |
Antispila treitschkiella | AAU1917 | DL-AT2 | HELA149–17 | Cornus mas | KY613516 | NHMUK |
Antispila treitschkiella | AAU1917 | DL-AT3 | HELA147–17 | Cornus mas | KY613517 | NHMUK |
Antispilina ludwigi | AAW5935 | RMNH.INS.11852 | HELA034–13 | Bistorta officinalis | MG191940 | RMNH |
Antispilina ludwigi | AAW5935 | RMNH.INS.11855 | HELA033–13 | Bistorta officinalis | MG191943 | RMNH |
Coptodisca lucifluella | AAV9339 | RMNH.INS.24368 | HELA047–13 | Juglans regia | KJ427013 | RMNH |
Coptodisca lucifluella | AAV9339 | RMNH.INS.24462 | COPIN003–14 | Juglans regia | KJ427007 | RMNH |
Heliozela hammoniella | AAK4551 | MM15506 | LEFIG642–10 | JQ412566 | ZMUO | |
Heliozela hammoniella | AAK4551 | MM15507 | LEFIG643–10 | HM876306 | ZMUO | |
Heliozela resplendella | AAI6721 | MM06188 | LEFID308–10 | HM873105 | ZMUO | |
Heliozela resplendella | AAI6721 | RMNH.INS.24359 | HELA058–13 | MG191938 | RMNH | |
Heliozela sericiella | AAK4533 | MM05954 | LEFIA1161–10 | JQ412568 | ZMUO | |
Heliozela sericiella | AAK4533 | MM05955 | LEFID174–10 | HM872981 | ZMUO | |
Heliozela sp. Lesbos | ACO3181 | RMNH.INS.24606 | HELA102–14 | MG191946 | RMNH | |
Holocacista rivillei | AAU2507 | AVS09 | ANTVI027–11 | Vitis vinifera | JQ412569 | UPI |
Holocacista rivillei | AAU2507 | RMNH.INS.24626 | HELA107–14 | Vitis vinifera cv Pinot Grigio | KP697786 | RMNH |
DNA extraction and sequencing. DNA was extracted using the Qiagen Blood and Tissue kit, either destructively from larvae or adult specimens preserved in 96% or 100% ethanol or non-destructively from the abdomen of voucher specimens, which were then used to prepare genitalic dissections (partly according to protocol in
Data selection and phylogenetic analysis. Since many molecular data on Heliozelidae have recently been published or will soon be published (
The sequence data generated and used in this study have been deposited in the public BOLD dataset (“Antispila treitschkiella and petryi in Europe ” [DS-ANTITR], doi: 10.5883/DS-ANTITR and GenBank (Table
Comparison of measurements of forewing and costal spot in Antispila petryi and A. treitschkiella males and females. Measured on photos, specimens from RMNH collection. Forewing length here does not include fringe, and thus differs from measurements in descriptions. The ratio length–width of the costal spot differs significantly between the species (marked in bold), but still shows an overlap.
Antispila petryi | Antispila treitschkiella | |||||
---|---|---|---|---|---|---|
Range | Mean ± sd | # | Range | Mean ± sd | # | |
Males | ||||||
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 | 1.179–1.625 | 1.373 ± 0.154 | 7 | 0.923–1.222 | 1.092 ± 0.096 | 11 |
Ratio spot w/fw l | 0.098–0.148 | 0.129 ± 0.019 | 7 | 0.116–0.157 | 0.135 ± 0.014 | 11 |
Females | ||||||
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 | 1.045–1.517 | 1.245 ± 0.165 | 7 | 0.780–1.171 | 0.974 ± 0.127 | 10 |
Ratio spot w/fw l | 0.110–0.167 | 0.132 ± 0.019 | 7 | 0.124–0.169 | 0.151 ± 0.013 | 10 |
Hostplant nomenclature and recognition. For the nomenclature we follow in principle
In Europe, there are three widespread native species of Cornus, two shrubs C. mas and C. sanguinea, and a herb, C. suecica L. In addition C. alba L. occurs natively in Russia, North and East of Moscow, through Siberia to Korea and NE China (
For C. sanguinea in Europe there usually are two subspecies recognised: C. sanguinea subsp. australis (C.A.Mey.) Jáv. from SE Europe (but often planted elsewhere!), and the nominal subspecies sanguinea. They can be separated by the type of hairs on leaf-underside: normal hairs in sanguinea, so-called compass hairs or medifixed hairs (also the common type of hairs in other Cornus spp.) in subsp. australis (see Suppl. material
In our experience, many records of hostplants of Antispila are either incomplete by just mentioning the genus Cornus, and quite a few are incorrect or questionable. We encourage collectors and observers to try to identify the host accurately to species level, in order to get a better impression of the host distribution of the Antispila species. Identification is usually not difficult when fruits are present, which may be found on the ground later in the season. Identification of single plucked leaves may be difficult, but often not impossible, and it is best to make notes and photographs and take a voucher (herbarium sample) of the plant where possible for future reference. As a host identification aid, we list some of the vegetative characters of the four species, and add the Japanese Cornelian Cherry, C. officinalis Torr. ex Dur., since A. treitschkiella mines can be found on botanic living collections in Europe seemingly misidentified as that species, or of hybrid origin:
Cornus mas. Stems green, rarely reddish. Leaves 4–10 cm long, greatest width below middle. Veins 3–5 pairs, often with tufts of white hairs in vein axils, most hairs medifixed, straight (Suppl. material
Cornus officinalis. Stems green or red. Leaves ovate to elliptic, 4–12 cm long. Veins 4–7 pairs, with dense tufts of rusty red to brown (not white) hairs in vein axils. Otherwise closely resembles C. mas. (Suppl. material
Cornus sanguinea subsp. sanguinea. Stems at least partly red. Leaves 3–10 cm long, largest width in middle. Veins 3–4 pairs, no tufts of hairs, but hairs spread on green underside (becoming red in autumn), only few medifixed hairs (Suppl. material
Cornus sanguinea subsp. australis. As nominal subspecies, but most hairs are medifixed (Suppl. material
Cornus sericea. Stems red at sun side, forming roots when touching ground (stoloniferous). Leaves 6–13 cm long. Veins 5–7 pairs, underside often hairy, pale coloured (glaucous), both sides with medifixed hairs (Suppl. material
Cornus alba. Stems always red to purple, rather straight, not forming roots when touching ground. Leaves 4–10 cm. Veins 5–7 pairs, underside often hairy, pale coloured (glaucous), both sides with medifixed hairs.
Living collections study. A few parks and living botanic collections in the UK were studied in detail in 2016–2017 in order to address the reliability of findings regarding monophagy of A. treitschkiella and A. petryi in particular. Because A. treitschkiella was found to be commonly established in the London area as well as in Cambridge, reaching outbreak levels on some trees of C. mas, and Cornus species were often planted in clusters, this seems a reasonable assay of hostplant repertoire at this time. This survey was conducted in the UK in October and November 2016 within the distribution range so far known for A. treitschkiella, guided mostly by garden maps printed by botanical garden staff. The survey was mainly conducted at Royal Botanic Gardens, Kew on 28.ix.2016 and at Royal Horticultural Society, Wisley on 5.x, with a visit to Cambridge Botanical Gardens on 17.ix. Depending on different plantings at these gardens, examination was conducted of C. mas and C. sanguinea (including several cultivars of both species), and in addition for specimens of C. sericea, C. rugosa Lam., C. alba, C. amomum Mill., C. brachypoda C.A. May, C. schindleri Wangerin, C. drummondi C.A. Mey, C. darvasica (Pojark.) Pilip., C. kousa F. Buerger ex Hance (incl. “Eddie’s White Wonder”, C. walteri Wangerin (=C. coreana Wangerin), C. controversa Hemsl., C. bretschneideri L. Henry, C. capitata Wall., C. × arnoldiana Rehder (= C. racemosa Lam. x C. obliqua Raf.), C. quinquinervis Franch. (= C. paucinervis Hance) and C. officinalis. A single specimen of C. mas was also examined at Chelsea Physic Garden in x.2016 and Oxford University Parks on 30.vii.2017, as well as a single specimen in Oxford Botanical Garden (C. mas var. variegata). Additional negative records on C. mas from other observers elsewhere in the UK are reported under results.
Herbarium study. In the UK in 2016, herbaria in Kew and Wisley and at NHMUK (the AMC herbarium of native and naturalised species) were examined and Cornus was surveyed, including all available C. mas, with the hope of determining origins of invasion of A. treitschkiella in the UK and within Europe. It was hoped that this approach would prove informative as it was for Cameraria ohridella Deschka & Dimič (
Parasitoids. We have tried to trace all literature records of parasitoids of these species. As basis we used the databases for Chalcidoidea (
Parasitoids recorded from Cornus feeding Antispila species. Where possible, primary sources were checked, indicated by the catalogue databases (
Taxa | Locality | Host-plant | Original identification | Reference |
---|---|---|---|---|
Antispila petryi | ||||
Eulophidae | ||||
Pnigalio pectinicornis (Linnaeus, 1758) | UK | C. sanguinea | A. treitschkiella |
|
Pnigalio cf. soemius (Walker, 1839) | UK, Surrey, Priest Hill, | C. sanguinea | A. petryi | DCL reared |
Antispila treitschkiella | ||||
Braconidae | ||||
Colastes flavitarsis (Thomson, 1892) | HU: Pécs, 17.xi.1971, 27.x.1974 | C. mas | A. treitschkiella |
|
Eulophidae | ||||
Chrysocharis budensis (Erdős, 1954) | HU: Budapest, Budakeszi, leg. Szőcs | [interpreted as C. mas, since Szőcs recognised both Antispila spp.] | A. treitschkeella |
|
Chrysocharis budensis (Erdős, 1954) | PL: Woj. Poznan | Not given | A. stachjanella |
|
Cirrospilus diallus Walker, 1838 | CZ: Koda near Beroun, 3.1954 | C. mas | A. sp |
|
Cirrospilus viticola (Rondani, 1877) [as subviolaceus Thomson] | CZ: Koda near Beroun, 3.1954 | C. mas | “a leafminer” [interpreted as treitschkiella] |
|
Cirrospilus vittatus Walker, 1838 | CZ | C. mas | A. sp |
|
Derostenus gemmeus (Westwood, 1833) | HU: Normafa, 14.vii.1954 | [interpreted as C. mas, since Szőcs recognised both Antispila spp.] | A. treitschkeella |
|
Minotetrastichus frontalis (Nees, 1834) | UK, SE London, Kelsey Park, x.2016 | C. mas | A. treitschkiella | DCL reared |
Minotetrastichus frontalis (Nees, 1834) | UK: London, Victoria Embankment, 30.vi.2017 | C. mas | A. treitschkiella | DCL reared |
Antispila metallella | ||||
Eulophidae | ||||
Chrysocharis budensis (Erdős, 1954) | NL: Simpelveld [as Simpelweld] | Not given | A. pfeifferella |
|
Chrysocharis budensis (Erdős, 1954) | PL: Torún | C. sanguinea | A. pfeifferella |
|
Host unidentifiable Antispila | ||||
Colastes braconius Haliday, 1833 | UK | Swida sanguinea | A. sp. |
|
Colastes braconius Haliday, 1833 | Not given | A. treitschkiella |
|
|
Colastes flavitarsis (Thomson, 1892) | HU: Pécs, Üszög,1.x.1973 | C. mas | A. pfeifferella [unlikely] |
|
Chrysocharis budensis (Erdős, 1954) | DE: München | Not given | A. treitschkiella |
|
Pnigalio cristatus Ratzeburg, 1848 | [primary source not seen] | Not given |
|
|
Zagrammosoma variegatum Masi, 1907 | [primary source not seen] | Not given |
|
Adult morphology.
Venation.
Genitalia.
In the female genitalia,
Larval characters. In the original description of A. petryi,
Mines. The shape of the mines is variable, and both earlier authors were unable to point out clear differences, other than the hostplant species. The size of the cut-out is on average slightly smaller in A. petryi than treitschkiella, but without diagnostic value (Figs
Heliozela Herrich-Schäffer, 1853
1. hammoniella Sorhagen, 1885
2. resplendella (Stainton, 1851)
3. lithargyrellum (Zeller, 1850)
4. sericiella (Haworth, 1828) [type species]
Antispila Hübner, 1825
5. metallella (Denis & Schiffermüller, 1775) [type species]
6. treitschkiella (Fischer von Röslerstamm, 1843)
7. petryi Martini, 1899
Provisionally in Antispila, but not belonging there
8. oinophylla van Nieukerken & Wagner, 2012*
ampelopsifoliella auctt. [misidentification]
Antispilina Hering, 1941
9. ludwigi Hering, 1941 [type species]
Holocacista Walsingham & Durrant, 1909
10. rivillei (Stainton, 1855) [type species]
Coptodisca Walsingham, 1895
11. lucifluella (Clemens, 1860)*
* species introduced from North America
1 | Forewing shiny grey to brown, with only one or two spots along dorsum | Heliozela [not further treated here] |
– | Forewing with pale spots at both costal and dorsal margin, often a fascia present as well | 2 |
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 Juglandaceae: walnut species (Juglans L.) and pecan (Carya illinoinensis (Wangenh.) K.Koch) in Italy ( |
Coptodisca lucifluella |
– | Forewing more or less uniform brown or fuscous, with a pale fascia, sometimes broken, at 1/3 and two opposite spots at 2/3 | 3 |
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 Vitaceae: grapevine (Vitis vinifera L.) and Virginia creeper (Parthenocissus quinquefolia (L.) Planch.) in Italy ( |
Antispila oinophylla |
– | Forewing without an additional apical spot, head not white, various tinges of grey, brown or leaden. Native species | 4 |
4 | Small species, wingspan 4.0–4.7 mm. Venation reduced, no cell in forewing. Antenna with 15–20 segments. Leafminers on Vitis or Bistorta Scop. | 5 |
– | Larger species, wingspan at least 5.0 mm, usually more. Venation complete with discoidal cell (Figs |
6 |
5 | Forewing fascia usually divided into two spots, spots relatively large, antenna with 15 segments. Male foretibia with small epiphysis. Mediterranean species, mines on Vitis ( |
Holocacista rivillei |
– | Forewing fascia not divided in two spots, spots small, antenna with 20 segments. Male foretibia without epiphysis. Rare Central European species, mines on Bistorta officinalis Delarbre (Polygonaceae) ( |
Antispilina ludwigi |
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 |
7 |
– | 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. | Antispila metallella |
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: C. sanguinea | A. petryi |
– | 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: C. mas | A. treitschkiella |
1 | Uncus medially blunt, central lobe reduced or absent. Valva at inner margin with spinelike process (Figs |
A. metallella |
– | Uncus with prominent central lobe. Valva without spinelike process | 2 |
2 | Horseshoe-shaped anellus present between the valvae (Fig. |
A. petryi |
– | No anellus between valvae visible (Fig. |
A. treitschkiella |
1 | Near start of mine 2–7 small brown scars made by ovipositing female (“test punctures”) (Figs |
A. metallella |
– | No scars near start of mine. Larva with row of dorsal black sclerotised plates (note: larvae usually feed with ventral side upwards) (Figs |
2 |
2 | Mine on C. mas. Full grown active larva whitish, with maximum of 8 black dorsal plates (Fig. |
A. treitschkiella |
– | Mine on C. sanguinea. Full grown active larva greyish, with maximum of 9 black dorsal plates, often less visible (plates not always melanised), plate on mesothorax present (Fig. |
A. petryi |
Antispila petryi Martini, 1899: 398. Syntypes: Germany: 12–14 adults, Thüringen, Sachsenburg, caterpillars 18.viii.–8.ix.1895, Cornus sanguinea, emerged 6–24.vi.1896, 1898, Martini; ditto, caterpillars mid ix.–3.x.1897, no emergence date given (at least seven syntypes in NHMUK via Hofmann collection, examined by DCL).
Antispila
petryi
;
Elachista
treitschkeella
[Misidentification, Unjustified emendation];
Elachista
treitschkiella
[Misidentification];
Antispila
treitschkiella
[Misidentification];
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
Differential diagnosis. Antispila petryi and A. treitschkiella differ from A. metallella by their smaller size (wingspan 4.8–7.0 mm against 6.8–8.5 mm) and in male by presence of a tuft of yellow androconial scales on forewing underside. A. petryi differs from A. treitschkiella by the usually smaller and more triangular costal spot at 2/3, and A. petryi is on average smaller than A. treitschkiella, but there is some overlap (wingspan 4.8–6.1 against 5.7–7.0 mm). In the male genitalia, the indentations in the uncus are shallower in A. petryi than in A. treitschkiella, the lateral process of the transtilla is straight and widened and the shorter phallus bears two types of spines externally. The horseshoe-shaped sclerotized anellus is characteristic for A. petryi, an anellus is undeveloped in A. treitschkiella. Separation by female genitalia not reliable.
Larva easily separated from A. metallella by presence of a row of dorsal black dots, in contrast to A. treitschkiella, a total of nine dots, including the mesothorax, but some of these may be poorly melanised, making this character not always useable; abdominal segment 8 with a single row of five black warts. Leafmines in principle not separable without larva or when hostplant species is not known.
Male (Figs
Antispila, adult habitus, all from Switzerland. 1 A. petryi, male, Mont Vully, Sugiez, e.l. 08.v.2016 2 A. petryi, female, Mont Vully, Sugiez, e.l. 30.iv.2016 3 A. treitschkiella, male, Gampelen, e.l. 17.iv.2016 4 A. treitschkiella, female, Le Landeron, e.l. 17.iv.2016 5 A. metallella, male, Muntelier, 30.iv.1990 6 A. metallella, female, Cudrefin, e.l. 18.iii.2015. All to same scale, scale line 10 mm. Photos: Rudolf Bryner.
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
Hostplants. Cornus sanguinea, both subsp. australis and sanguinea, and incidentally on cultivated C. alba (Wocke cited in
Leafmines (Figs
Life history. Univoltine. Larvae usually from late August until October, in Greece still active in early November, few records from early August. Adults emerge in captivity from April to June, the few specimens collected as adults being found from June to early August. We assume they are mostly active during the day, rarely collected at light, but found in malaise traps.
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 A. treitschkiella, but correct records (on the basis of examined adults, larvae or hostplant data) exist from Germany, Poland, Hungary, Czech Republic (Z. Laštůvka, pers. comm.), Austria (as A. treitschkiella:
The natural distribution of C. sanguinea comprises most of Europe, in the north including the whole of the British Isles, southern coastal areas of Norway and Sweden (south of Stockholm), a northern limit in Estonia and in Russia below a line from the Latvian/Estonian border to Moscow, in southern Europe including northern parts of the Iberian peninsula, all of Italy, Corsica, Sardinia, and all of the Balkans except the Greek islands; local in northern Turkey, and widespread in the Caucasus region, reaching Iran (
This species was described from an unspecified number of specimens reared by Martini from mines on C. sanguinea, collected as caterpillars in 1895 and 1897 in Sachsenburg (not far from Sömmerda) and with adults reared in 1896 and 1898. Martini also mentioned the species from Regensburg, Bavaria, and Breslau (now Wrocław, Poland), but he probably did not study these himself, and relied on information received from respectively Hofmann and Wocke. Issue no 10–12 (Heft IV), pages 333–429 of volume 59 of the Stettiner Entomologische Zeitschrift was published in June 1899 (see page 429: http://biodiversitylibrary.org/page/8946769). This would have allowed Martini to include the reared adults from 1898, although he did not give emergence dates for these.
Martini’s study is very thorough, comparing A. petryi with A. treitschkiella and A. metallella (“pfeifferella”), including the biology and mines. Martini’s collection of Palaearctic Microlepidoptera was presented to the primary school (“Volksschule”) in Sömmerda, his place of residence in the German federal state of Thüringen, and is no longer traceable (
Note. This species was known under the three names listed under the synonymy [viz. treitschkiella, petryi and stachjanella].
The male genitalia figured by Wojtusiak and reproduced by Razowski resemble more A. petryi than A. treitschkiella, but some important characters are not illustrated (transtilla, anellus). Also in the drawings in
Elachista treitschkiella Fischer von Röslerstamm, 1843 [March]: 297, pl. 100: 4. Syntypes, number and sex unspecified: Austria: Wien [Vienna], v–vi, leg. Mann, flying around Cornus mascula. One potential syntype in NHMUK without date or hostplant information [examined by DCL; not dissected].
Oecophora treitschkiella Duponchel, [1843, 4 May]: 319, pl. 77:1. Syntypes, [Austria: Vienna, leg. Mann], received from Pareyss (independent new description) (Paris).
Antispila
stachjanella
;
Antispila
treitschkiella
;
Antispila
treitschkeella
[unjustified emendation]; [
Antispila
treitschkella
[misspelling];
Antispila
stachjanella
;
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 A. petryi. Larva see A. petryi, characteristic absence of black dot on mesothorax (second visible segment), making a maximum of eight dots, abdominal segment 8 with 2–3 rows comprising many black warts of different sizes.
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
Antispila petryi (27, 29, 31) and A. treitschkiella (28, 30, 32), comparative details of male and female genitalia. 27, 28 Uncus, slides JCK8056, JCK7888 29, 30 Phallus, phallotheca and juxta, slides JCK8056, JCK8059 31, 32 Tip of oviscapt, slides D1258, D1257 (D=Doets). Drawings Sjaak Koster.
Female genitalia (Figs
Larva (Figs
Antispila species, details of larvae in microscopic slides, stained with phenosafranin, Switzerland. 33–35 A. petryi, Le Landeron, RMNH.INS.30614 and Mont Vully, RMNH.INS.30615 (34) 36–38 A. treitschkiella, Le Landeron, RMNH.INS.30611 and Gampelen, RMNH.INS.30613 (38). Scales 1 mm (larvae), 200 μm (heads), 100 μm (abdominal tips). Photos Erik van Nieukerken.
Antispila species, leaves of Cornus with vacated leafmines, dry collection RMNH. 55 C. sanguinea with A. petryi, RMNH.INS.41141, Netherlands, Limburg, Geulhem, 26.viii.1949, C. Doets 56 C. mas with A. treitschkiella, RMNH.INS.40829, Netherlands, Zuid-Holland, Leiden, 17.x.2004, E.J. van Nieukerken. Photos Erik van Nieukerken.
Hostplants Cornus mas. Records on C. sanguinea probably all refer to A. petryi.
The larvae are monophagous and do not occur on the shrubs ofCornus sanguineaL. which species is also very frequent in the forest examined.
There are some records on the planted species C. sericea or C. alba, but the accuracy of these is questionable. May be occasionally also on C. officinalis (see below under Living collections).
Leafmines (Figs
Life history. Bivoltine. Larvae of the first generation are found from June to July, a second generation from August until early November; it is possible that generations may overlap. Adults emerge in captivity from April to June (the few earlier records are almost certainly forced breedings), and again from July to late August; the few records of field caught adults agree with this pattern. The adults may swarm around the host during the day, as already was mentioned by Fischer von Röslerstamm (1843). A recent observation of ca. 100 swarming adults on C. mas in the Netherlands can be seen here: https://waarneming.nl/waarneming/view/139225815.
Widespread in central and southern Europe, in the natural range of C. mas, and north and west of it occurring on the frequently planted trees. Positively recorded on basis of adult or larval characters, or hostplant: England, the Netherlands, Belgium, France, Germany, Poland, Czech republic, Slovakia, Hungary, Switzerland, Austria, Italy, Slovenia, Bulgaria, and Ukraine. We record it here from Albania, one specimen on BOLD, GRPAL724–11, agreeing in DNA barcode (Table
The natural distribution of C. mas is much more restricted than that of C. sanguinea, its NW border being from SE Belgium to NW France, away from the coast, covering large parts of France, whereas it is scarce in isolated regions in Germany, West and South Switzerland, Austria north and east of the Alps, large parts of Italy and the whole of Southeast Europe south of southern and western Czechia, Slovakia, and southern parts of the Ukraine; also coastal areas of Turkey, Caucasus region and Black Sea coast of Russia and Crimea, just reaching Azerbaijan and North Iran. Cornus mas is absent from the Mediterranean islands and the Iberian Peninsula, except for a very small area in the Catalonian Pyrenees (Da Ronch et al. 2016).
The original description by Fischer von Röslerstamm (1843) of Elachista treitschkiella is based on an unspecified number of specimens, collected by the renowned Austrian collector J. Mann in the vicinity of Vienna, where the adults were observed flying in the afternoon and particularly swarming around sunset about bushes of “Kornelkirschensträucher (Cornus mascula)”, i.e. the European cornel or C. mas. The hostplant and the detailed description leave little doubt as to the identity of this moth, also in the light of our DNA barcode findings. Fischer von Röslerstamm named this species after Friedrich Treitschke, who died a year earlier on 4.vi.1842, and he added a long obituary in a footnote. Duponchel [1843] described the species independently again, as Oecophora treitschkiella, also based on material collected by Mann, most likely in Vienna. De
Fischer von Röslerstamm’s Microlepidoptera were acquired in 1843 by Herrich-Schäffer (
Antispila stachjanella was described from an unspecified number of specimens of both sexes, bred by Dziurzyński in 1944–1947 [1948] from larvae mining the leaves of C. mas in Kraków. In the Polish Academy of Sciences, Kraków are about 150 (undissected) specimens of A. stachjanella from Dziurzyński’s rearings (1944–1948), none of which is clearly labelled as any kind of type, although at least some of them constitute a potential syntypic series (L. Przybyłowicz, pers. comm.). In NHMUK there are four specimens under A. stachjanella, two of which on loan, and two of which are labelled “COTYPUS”, only one of which (NHMUK010305235) is apparently in the correct date range (the other NHMUK010305235, is labelled 1949). The original description of A. stachjanella is a short diagnosis in Latin amidst a long Polish text and a rather long English summary of six pages. The Latin text does not give information on types, but there is a drawing embedded on page 5, showing a male and a wing of a female collected in Kraków Botanical Garden which emerged 26.vi.1948 ex larva (Dziurzyński 1948: 5). In Krakow there are 11 corresponding specimens which are undissected, two males and nine females (L. Przybyłowicz, pers. comm.), one of which would potentially be suitable for lectotype designation. The whereabouts of specimens giving rise to illustrations of genitalia are unknown. Both in the collections in Leiden and London there are four specimens each of A. treitschkiella from Kraków, collected by S. Błeszyński between 1946 and 1948. In Kraków there are no specimens currently labelled A. petryi and 36 specimens under A. treitschkiella 14 of which collected by S. Błeszyński up until 1948. Błeszyński is acknowledged by
It is somewhat mysterious why Dziurzyński needed to introduce A. stachjanella in such a detailed study, while he only briefly addressed the separation from A. treitschkiella. A single specimen identified by Hering and collected by Toll in Podolia was his only comparative material, and he based the different identities on the fact that the antennae of his species were ringed, whereas the specimen of A. treitschkiella he studied did not have rings. He also cited
Antispila treitschkiella has been – under the name A. stachjanella – the subject, of two very detailed studies respectively on larval behaviour and the central nervous system (
Tinea metallella [Denis & Schiffermüller], 1775: 144 (Dunkelgoldener Schabe mit 6. Silberzeichen). Syntypes, unspecified: Austria: Wien [Vienna] [not examined, collection lost by fire]
Tinea
pfeifferella
Hübner, [1813]: pl. Tineae IV, Nobiles, 59: fig 398. Syntypes, unspecified, Europe [not examined]. (Lost). A primary homonym of Tinea pfeifferella Hübner, [1813]: pl. 63: fig. 422 (see
Antispila stadtmuellerella Hübner, [1825]: 419. Syntypes, unspecified, Europe (Hübner) (Lost). [Replacement name for the pre-occupied Tinea pfeifferella, type species of the genus Antispila Hübner, 1825]
Tinea
quadriguttella
Haworth, 1828: 574. Synonymised by
Elachista
pfeifferella
;
Antispila
pfeifferella
;
Antispila
metallella
;
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
Differential diagnosis. Adults (Figs
(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.
Larva. A detailed description of a 4th instar larva was given by
Antispila metallella, leafmines and larvae. 62 Leafmine with larva, Netherlands, Oegstgeest, on Cornus sericea, larva RMNH.INS.18464. 63 Vacated leafmine on C. sanguinea, Netherlands, Valkenburg, ZMA.INS.MIG.02122. 64 Larva and halves of shield, larva RMNH.INS.18464. 65 Vacated leafmine on C. sanguinea, Bulgaria, Ilidentsi, RMNH.INS.42092. Scales 5 mm. Arrows indicate test punctures. Photos Erik van Nieukerken.
Antispila treitschkiella, invasion in the UK and herbarium specimens. 66 Leafmines of on Cornus mas at London, Hyde Park Corner, 22.ix.2016. 67 Leafmines on C. mas at London, Temple, Victoria Embankment, 28.ix.2016. 68 Herbarium specimen, RBG Kew, Verdcourt 4160, 22.viii.1964, from Mt. Pelion, Greece: C. mas with mines of A. treitschkiella 69 Detail of 68 70 Herbarium specimen, RBG Kew, Schneider 1681, 28.viii.1907, from Rila Mt., near Samakov, Bulgaria, C. mas with mines of A. treitschkiella. Photos David Lees.
Hostplants Cornus sanguinea, both subsp. australis and sanguinea, rarely also recorded from planted C. alba and C. sericea (see herbarium results). In literature repeatedly recorded from C. mas, but in most cases without any data nor references: these records are unlikely and should be checked. Some leafmine material of A. metallella seen by us and labelled as from C. mas appeared to be misidentified, either the insect (being A. treitschkiella) or the plant (being C. sanguinea).
Leafmines. Leafmines are larger than those of the other two species, and the species can be separated by the presence of test punctures near the oviposition site (Figs
Life history. Univoltine. Larvae from early June to early August, much earlier than A. petryi on the same hostplant. Adults fly from April to early June.
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 Tinea pfeifferella in the same work for two different species; it was his decision as first reviser to replace the name published on the earlier plate (pl. 59) rather than the one on the later (pl. 63) with “Antispila Stadtmüllerella” (
We included all ten European species of Heliozelidae (39 specimens) and one unnamed candidate species in the DNA barcoding analysis. The barcodes of all group as monophyletic clusters and have much larger interspecific than intraspecific distances (Fig.
Identifying material based on DNA barcodes is highly reliable for this group and provides an alternative to morphology. True A. treitschkiella belong to the cluster with Barcode Index Number (BIN) BOLD: AAU1917 and A. petryi to BOLD: AAV5055, while A. metallella belong to BOLD: ACG8679.
About 20 species or cultivars of Cornus were examined among living collections in the UK (see Methods). Every single C. mas tree examined by DCL had mines of A. treitschkiella, except for C. mas and C. mas “var. variegata” at Oxford Botanic Garden (where A. treitschkiella still seems to be absent). Perhaps surprisingly, neither A. petryi nor A. metallella were positively identified in the botanic gardens living collections survey, although it is possible that old mines on C. sanguinea “Annie’s Winter Orange” at Wisley represented the former and on “C. alba” at Kew represented the latter. A. petryi was present on C. sanguinea at Priest Hill Reserve, Surrey, and along hedgerows at other countryside locations in the Cambridge area, and old mines probably of this species were present on C. sanguinea in Kensington Gardens, London.
At Kew and Wisley, living specimens labelled as C. paucinervis (a synonym of C. quinquinervis Franch.) with mines of A. treitschkiella turned out to be misidentified and were in fact referable to C. mas (K. McGinn, pers. comm.). The same applied to specimens labelled as C. officinalis (a species in the same subgenus as C. mas), with one exception: a tree of this species at Wisley with not a single mine of A. treitschkiella had the character of typical C. officinalis, dense rusty red hairs in the axils of the underside leaf venation. In the Wisley shop, the few living specimens for sale labelled as C. officinalis (without rusty hairs in the leaf axils) had one to two mines per plant; we therefore regard this record as inconclusive. There are also three specimens of A. treitschkiella (as A. stachjanella) that A. Dziurzyński reared in 1946–1949 from C. officinalis from the botanic garden in Kraków, in the collection of the Polish Academy of Sciences. The identity of any so-labelled plants still existing in that botanic garden would also need to be verified.
At Cambridge Botanical Garden in October 2016, all C. mas specimens had strong prevalence of A. treitschkiella mines (usually at least 1–10 per branch). In one case, an adjacent specimen of C. amomum subsp. obliqua (Raf.) J.S. Wilson (labelled as “C. obliqua”; number 37.0114) had a few mature mines, apparently as spillover from an adjacent C. mas. As these few moth individuals have not yet been DNA barcoded, nor the plant identity rechecked by botanists, we regard also this potential new hostplant record for A. treitschkiella so far as also inconclusive. This Cornus species is among the North American Kraniopsis Raf. (subgenus) and is either a subspecies or a good species related to C. amomum (
As C. officinalis is the inferred sister to C. mas (
In Wisley, the 14 specimens of C. mas at a suitable stage of leaf maturity collected between 1848–1997, evidenced zero mines of A. treitschkiella. Unfortunately, there were relatively few cultivated collections of C. mas preserved since the 1980s.
At NHMUK (AMC herbarium), the handful of specimens of C. mas of UK origin were also clear of mines. However, in this last collection, two specimens of naturalised C. sericea (labelled as C. stolonifera) from Panshanger, Hertfordshire, 1952, had clear mines of A. metallella, and the mature larva was extracted for verification in one case. This confirms one of the previous hostplant records for A. metallella; the species is, however, also reported from C. alba (
At Kew Gardens herbarium, only four specimens of C. mas were found with mines apparently of A. treitschkiella, all from native collections in continental Europe (Greece: Mt. Pelion; Albania: N. of Merkopele; France: Charance; Bulgaria: Rila Mts, Samakov; for details see Suppl. material
The herbarium of the Naturalis Biodiversity Center, Leiden was briefly scanned by means of its BioPortal (http://bioportal.naturalis.nl/), but not a single sign of mines was found so far.
Our results show that the C. sanguinea feeding A. petryi differs in many aspects from the C. mas feeding A. treitschkiella, and thus we conclude that the synonymy by Razowski (1978) was unwarranted. Identification is straightforward on the basis of male genitalia, larval characters, hostplant and DNA barcodes. For a separation of A. stachjanella from A. treitschkiella we, however, do not find any support, and there is thus only one taxon feeding on C. mas.
Literature and other records of Antispila “treitschkiella” that specifically provide the hostplant species name can be attributed to either A. petryi or A. treitschkiella s. str., but many records are inconclusive in this regard.
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 Cornus species have a different leafminer fauna is no surprise, as the species are not closely related. Cornus mas belongs to the subgenus Cornus L. that comprises a few Asian and one European species, and is easily recognised by the cauliflory, yellow flowers appearing before the leaves, and edible large red fruits, resembling a cherry (hence the name Cornelian cherry). On the other hand, C. sanguinea belongs to the large subgenus Kraniopsis Raf., with numerous Asian and North American species: these have white flowers in umbels, appearing after the leaves, and small white or blue berries. Phylogenetically, both subgenera are clearly separate (Xiang et al. 2006). Cornus alba and C. sericea also belong to Kraniopsis. Up to now, however, we have not seen evidence for A. petryi on these hosts and only record here A. metallella from C. sericea.
In summary, all our results support the hypothesis that the two species A. treitschkiella and A. petryi are monophagous, respectively on C. mas and C. sanguinea. In an interesting experiment,
The first reports of a range expansion of A. treitschkiella on to planted C. mas was by
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 C. sanguinea. Large numbers of mines were found on C. mas in a few localities in the provinces of Gelderland and Noord-Brabant, but in many places no mines were present. It is now clear that these records represented the first Dutch records of the real A. treitschkiella, that since then has become numerous in many places throughout the Netherlands (
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 A. treitschkiella collected on 23.viii.2016 in the South Kensington area of London. Since Antispila are not usually recorded in light traps, it may take some effort to detect any earlier records of adults in collections. After the initial finding, we observed mines on all examined trees verifiable as C. mas throughout London and beyond within the presently known range of A. treitschkiella, in London sometimes with very heavy impact. The UK herbarium survey proved negative, based however on few recent pressings of cultivated C. mas (the largest such collection being 14 specimens in leaf at Wisley from 1848–1997). Absence of evidence in this case does not constitute evidence of absence, except that it was possible for botanists, at least up to about 1997, to find sections of plants fully clear of mines. Cultivated C. mas are not sampled every year and even European collections from its natural range are rather sparse. At Kew and NHMUK, those few preserved specimens from the wild range add, however, to distributional knowledge, assuming these mines do not represent a species other than A. treitschkiella. The presence of the species on even decades-old stands of C. mas (e.g. at Priest Hill, Surrey) in the UK, that are probably separated from other populations by many kilometres suggests either long establishment or, more likely, surprising powers of adult dispersal aided by winds or road traffic and outbreak swarmings next to major roads (the last explanation has been invoked for rapid spread of Cameraria ohridella Deschka & Dimić: e.g.
We have no indication of expansion of A. petryi, even though its hostplant is also widely planted in parks and gardens, as are the Asian and American Cornus relatives. However, C. sanguinea has a much wider natural distribution than C. mas (
In the Netherlands A. petryi is only known to occur in the hills of South Limburg; it is absent from C. sanguinea at other sites where the host is native or where it is planted in parks and gardens. In contrast, A. metallella, feeding on the same host, has spread all over the country since the 1980s and is now common on planted C. sanguinea and sometimes is found on cultivated C. sericea and/or C. alba.
While many species have been observed in recent years to spread over Europe, whether they are native in Europe or not, the expansion of A. treitschkiella has remained partly unnoticed, as a result of some poor taxonomical decisions. It remains a mystery as to why the very extensive and detailed descriptions by
The expansion of A. treitschkiella to the north and west of the native range of its host on to planted trees fits the pattern observed in several other leafminers native to Europe. Especially species feeding on various Acer L. species (Sapindaceae) have been spreading in recent decades, such as Stigmella aceris (Frey) (Nepticulidae) on Acer campestre L. and A. platanoides L. (van
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 Cornus feeding species and we hope that the on-going expansion of these species will be followed by others in more detail. Posting photos of mines with accurate host data on observation sites such as https://observation.org/ is a useful way to make your records known.
Many people helped us with information or material, we are grateful for their help: Willem Ellis collected additional A. treitschkiella for DNA barcoding, Jurriën van Deijk kindly provided data from the Dutch Lepidoptera database Noctua. Bengt Å. Bengtsson, provided information on the population on Öland, and Urmas Jürivete on the Estonian population on Saaremaa. Carlos Lopez-Vaamonde kindly provided the barcode of a specimen from Albania, and BOLD staff is acknowledged for support in many ways. In the UK, Ian Kitching suggested the long established site of Priest Hill for a plantation of C. mas. Natalie Dale-Skey, Christer Hansson and John Noyes kindly assisted with identification of parasitoids reared from C. mas leafmines in the UK and Cees van Achterberg helped accessing literature data. Yvette Harvey and Andrew Salisbury at Wisley, Martyn Dickson at Edinburgh Botanic Garden and Kevin McGinn and Alex Monro at RBG Kew and at NHMUK (AMC collection) assisted with access to herbarium and living collections at Kew as well as Estela Dukan for the Sibbold Physik Garden. John Medhurst helped with suggesting localities for C. mas in London. Jozef Razowski and Lucasz Przybyłowicz kindly checked for potential type material of A. stachjanella in Warsaw and Sabine Gaal-Haszler searched for Elachista treitschkiella in Wien. Ruben Vijverberg and Frank Stokvis (Naturalis) are acknowledged for assistance with molecular lab work. The Naturalis library staff is acknowledged for continuous supporting literature requests. We thank reviewers Lauri Kaila and Zdenek Laštůvka and editor Maria Heikkilä for their insightful comments.
Specimen and Locality Data Antispila.
Data type: Excel file
Explanation note: Specimen data.
Figure S1. Cornus species, details of leaf underside. a C. sanguinea subsp. sanguinea, fresh leaf, Netherlands, Leiden, 26.ix.2017. b C. sanguinea subsp. australis, dried leaf, Greece, RMNH.INS.41827. c C. mas, fresh leaf, Netherlands, Leiden, 26.ix.2017. d C. sericea, dried leaf, Netherlands, RMNH.INS.41493. e C. officinalis, dried leaf, UK, Wisley 1996, Wisley herbarium.
Figure S2. NJ tree of barcodes, from BOLD.
Data type: 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).