2020, Entomologist’s Gazette 71: 221–247 doi: 10.31184/G00138894.714.1793 © Pemberley Books

Lyonetia prunifoliella (Hübner, 1796) (: ): a review of the species in England and consideration of L. padifoliella (Hübner, [1813])

R. J. HECKFORD1. Department of Life Sciences, Division of , Natural History Museum, Cromwell Road, London SW7 5BD, U.K.

S. D. BEAVAN The Hayes, Zeal Monachorum, Devon EX17 6DF, U.K.

Abstract This paper reviews British records of prunifoliella (Hübner, 1796) up to the end of 2017: all are from England. The account of the species in The and of Great Britain and Ireland 2 is considered and certain changes are suggested. Detailed descriptions of the , larval mine and pupa are provided, with illustrations. Larval foodplants are reviewed both of this species and L. padifoliella (Hübner, [1813]) as well as whether the latter has occurred in England. Key words: Lepidoptera, Lyonetiidae, , history in Britain, larval description, pupal description, foodplants, Lyonetia padifoliella

Introduction This paper was prompted by the discovery of a considerable number of mines of Lyonetia prunifoliella (Hübner, 1796) on spinosa L. in Devon, England, in the autumn of 2017. The species is accorded pRDB1 status in Great Britain by Davis (2012: 9), meaning that for the period 1980–2011 it had been recorded from five or fewer 10 km squares. Had Davis viewed it as an immigrant it would have been noted as such and not ascribed a status based on records from 10 km squares. All records from Great Britain for that period as well as records prior to and subsequent to the period 1980–2011 are from England. Prior to 2017 there are three records from the nineteenth century and the species was apparently not found in the twentieth century. A record from Worcestershire published in 1901 is a misidentification of (Linnaeus, 1758). A vacated mine on aucuparia L. found near Anstey, Leicestershire, on 27 July 1974, initially thought to be of this species, is a misidentification of sorbi (Stainton, 1861). The first twenty-first century records were in 2007, at two sites. Thereafter, prior to 2017 it has been noted on several occasions at different localities, and, with one exception, as adults and usually at light. As far as we are aware, before 2017 there have only been two records of larvae in Great Britain. The first was in 1893 in two gardens at Worthing, Sussex, on Malus sp. and Prunus japonica (sinensis) or Prunus sinensis. We consider what species was meant by these two Prunus names later in this paper and why we do not cite authors’ names for them in this section. The second occasion was of one

1. Scientific Associate. Correspondence address: 67 Newnham Road, Plympton, Plymouth, Devon PL7 4AW, U.K. 221 222 Entomologist’s Gazette (2020) Vol. 71 larva on in Kent in 2016. In late September 2017 in Devon the authors found over 340 mines of which just over 40 were tenanted, all on Prunus spinosa. As a result of informing others of this, numerous mines, most vacated, were found in Dorset, again all on Prunus spinosa. Details of these records are provided later in this paper. Lyonetia prunifoliella has two main forms. The typical one has a mainly white forewing with brown oblique dorsal and costal streaks, the first dorsal streak usually joining the first costal (‘the white form’), whereas f. albella (Eversmann, 1844) has a mainly brown forewing with a white, slightly sinuous, dorsal area to about one-half from which arises a white oblique streak to about the middle of the forewing and another white oblique streak of similar length from about two-fifths. Occasionally moths are found which are almost intermediate between these two forms. We are aware of one British county website that states that the species is sexually dimorphic. This is not correct. Both sexes have these forms. In Japan the two main forms have been viewed as seasonal, with f. ringoniella Matsumura, 1931, the one having a mainly white forewing, being called the spring form and f. malinella Matsumura, 1907, which appears to be the same as f. albella, being called the autumnal form (Kuroko, 1964: 24–26, pl. 1 figs 5a and 5b). We reared both main forms as well as intermediate examples from the larvae that we found in the autumn of 2017. The species has been increasingly recorded in England since 2017. It is beyond the scope of this paper to consider these subsequent records in any detail except to note that as well as adults larval mines have been found on various occasions. As far as we are aware all have been on Prunus spinosa. In mainland another similar species occurs, L. padifoliella (Hübner, [1813]). We consider both in the next section and later whether any English records might be of L. padifoliella.

Consideration of Lyonetia prunifoliella and L. padifoliella Like Lyonetia prunifoliella, L. padifoliella has two main forms. One has a forewing that is similar to but not exactly the same as that of the white form of L. prunifoliella in having a mainly white forewing with brown dorsal and costal streaks, but in this form of L. padifoliella the streaks are often darker brown and the first dorsal is far more oblique and does not join the first costal, and there is a narrow straight dark brown line extending from near its tip towards the termen. The other form of L. padifoliella has slightly similar markings to L. prunifoliella f. albella but usually most of the forewing is white or greyish white with two oblique dark brown dorsal streaks, although sometimes the forewing is more suffused generally with dark brown. Both species have short strigulae in the apical and tornal areas, often with a pale reddish brown area between them, a small black subapical spot, sometimes suffused with the ground colour in dark specimens, and a dark brown or black pencil projecting from the terminal cilia. In order to illustrate the morphological differences between the two species, Figs 1–4 show the white form of Lyonetia prunifoliella, L. prunifoliella f. albella and intermediates, and Figs 5–8 the white and brown forms of L. padifoliella. Figs 1–4 are moths reared by the authors from Prunus spinosa in England. Figs 5–6 are moths reared by R. J. D. I. Voith from Cotoneaster scandinavicus B. Hylmö in Norway and Figs 7–8 are moths reared by L. Aarvik from the same foodplant also in Norway. Entomologist’s Gazette (2020) Vol. 71 223

Hübner (1796–[1836]) named L. prunifoliella and L. padifoliella on different dates, both in Sammlung europäischer Schmetterlinge. The first was Tinea prunifoliella which is illustrated on plate 28, figure 191 in Part 8, published in 1796. Tinea padifoliella is illustrated on plate 46, figure 316 also in Part 8 but published between an unknown date in 1810 and 20 June 1813. We follow Hemming (1937: 212–213, 295 and 298) in ascribing the dates to these plates. Hübner’s coloured illustrations are small. Figure 191 on plate 28 (Tinea prunifoliella) shows a moth whose forewings are mainly white, with brownish or blackish streaks towards the apex. Figure 316 on plate 46 (Tinea padifoliella) shows a moth whose forewings are whitish in the costal area but with brownish or blackish streaks arising from the dorsal area. Stainton (1848: 2160) was the first to add L. prunifoliella to the British list. This was within ‘A Monograph on British Argyromiges’ and he gave it the specific epithet of padifoliella. Nevertheless, as shown in the next section, it is clear that he was referring to L. prunifoliella f. albella. However, it was the name padifoliella that he consistently used thereafter: Stainton (1852: 65; 1854a: 284; 1854b: 161; 1856: 58; [1857]: 112; 1859: 421). Subsequently, in most of the British literature this name has been placed as a junior synonym of L. prunifoliella, for example, South (1884: 39), Kloet & Hincks (1945: 138; 1972: 8), Emmet (1985: 224) and Agassiz, Beavan & Heckford (2013: 20), although Tutt (1898: 199) used the specific epithet padifoliella. Bengtsson & Johansson (2011: 421–423), however, show that L. padifoliella is a good species, separable both on forewing pattern and genitalia. They provide figures of both main forms of the adult L. prunifoliella but only the dark form of L. padifoliella, as well as the genitalia of both sexes of both species. Emmet (1985: 224), in The Moths and Butterflies of Great Britain and Ireland 2 (MBGBI), gives the most comprehensive account of L. prunifoliella in the British literature but not all the information provided can have been derived from English records, especially the larval foodplants and larval period given. This is unsurprising as Emmet would not have observed the species in England, since prior to 1985 there were only three records and all were from the nineteenth century. As some of the information provided by Emmet was almost certainly derived from mainland European sources his account may comprise both species. We consider this in a later section. Because the two species have been confused until fairly recently, this paper considers both species, including larval foodplants, and discusses whether any English records of L. prunifoliella might be those of L. padifoliella. We also consider whether L. prunifoliella is a very rare, or at least very local, resident in England or whether it is an immigrant that is an occasional temporary colonist.

English records of Lyonetia prunifoliella prior to the publication of The Moths and Butterflies of Great Britain and Ireland 2 in 1985 Up to and including 1985 Lyonetia prunifoliella had been recorded from only five localities in England: Whittlebury Forest, Northamptonshire (by 1848), near Stony Stratford, Buckinghamshire (by 1859), Boughton, Worcestershire (in 1869, but not published until 1901), Worthing, Sussex (in 1893, in two gardens), and near Anstey, Leicestershire (1974). Of these five, however, the Boughton record is a misidentification of L. clerkella and that from near Anstey was of one 224 Entomologist’s Gazette (2020) Vol. 71

Photo: Miss S. D. Beavan & R. J. Heckford Fig. 1. Lyonetia prunifoliella (Hübner, 1796), white form. England: Devon, Heybrook Bay, larva found 4.x.2017, moth emerged 24.x.2017.

Photo: Miss S. D. Beavan & R. J. Heckford Fig. 2. Lyonetia prunifoliella (Hübner, 1796) f. albella (Eversmann, 1844). England: Devon, Exminster, larva found 19.ix.2017, moth emerged 1.x.2017. Entomologist’s Gazette (2020) Vol. 71 225

Photo: Miss S. D. Beavan & R. J. Heckford Fig. 3. Lyonetia prunifoliella (Hübner, 1796) intermediate form. England: Devon, Exminster, larva found 22.ix.2017, moth emerged 4.x.2017.

Photo: Miss S. D. Beavan & R. J. Heckford Fig. 4. Lyonetia prunifoliella (Hübner, 1796) intermediate form. England: Devon, Exminster, larva found 22.ix.2017, moth emerged 5.x.2017. 226 Entomologist’s Gazette (2020) Vol. 71 -mine on which in fact proved to be that of Stigmella sorbi. We now examine all these in detail including those where the specific epithet has been given as ‘padifoliella’. In a later section we consider whether they are of that species. The first record was in a paper by Stainton (1848) on British Argyromiges Curtis, 1829. At page 2160 appears the following: ‘Sp. 32. PADIFOLIELLA, Hubner [sic] (fig. 30). Tinea padifoliella, Hubner [sic], Ti. 316. Lyonetia padifoliella, Zeller, Isis, 1839, 216.’ There then follows a detailed description of the adult. This certainly does not describe the white form of L. prunifoliella, nor the white form of L. padifoliella, but in our view it could represent the dark form of either species. Fortunately three plates accompany Stainton’s paper, which was published in two parts. All the plates have monochrome drawings by Stainton of the left hand forewing of the 40 species that are dealt with in his paper. Figure 30 (Fig. 9 in this paper) clearly shows Lyonetia prunifoliella f. albella. The only information that Stainton gives about its British occurrence is that it is ‘A local species, taken by Mr. Desvignes, in September, in Whittlebury Forest [Northamptonshire], off maple and whitethorn.’ ‘Maple’ is Acer campestre L. and ‘whitethorn’ is a Crataegus sp. Two years later Curtis (1850: 118) described Desvignes’ material as a new species, ‘A. [Argyromyges] Acerfoliella, Curt.; Padifoliella, Stain.’. He gives a detailed description and states: ‘For a pair of this rarity I am indebted to Mr. T. Desvignes, who took several in September and October flying out of maples and whitethorns in Whittlebury Forest. The sexes seem to vary considerably, but neither of them agrees with Hübner’s figure of T. Padifoliella, pl. 46. f. 316, in which the costa is white and the interior margin spotted dark, whereas in our species it is exactly the reverse.’ Whittlebury Forest remained the only known British locality until Stainton (1859: 424) recorded the species, still under the name Lyonetia padifoliella, from ‘Near Stony Stratford’ in Buckinghamshire, but gave no other locality. Quite why he did not include Whittlebury Forest is unknown to us. We have not been able to trace any more information about this record, which is repeated by Barrett (1905: 106), under the same name, who states ‘Stony Stratford—the only British locality given by Mr. Stainton’. Although it was the only locality given by Stainton in 1859, he had already added L. prunifoliella to the British list from Whittlebury Forest. This was an uncharacteristic oversight by Barrett. The third was from Boughton, Worcestershire in 1869, but not published until 1901 (Rea & Fletcher, 1901: 120). The only information provided is ‘Boughton (Fletcher)’. The Fletcher in this publication is not the W. H. B. Fletcher cited later in this section but J. E. Fletcher. Harper & Simpson (2003: 49), in The smaller Moths of Herefordshire & Worcestershire, state: ‘Lyonetia prunifoliella (Hübner) was recorded from Boughton, Worcester in 9/1869 but a specimen with the same date in Worcester Museum labelled prunifoliella has been examined and appears to be only an exceptionally large clerkella (Linnaeus), and therefore this record has been omitted as probably incorrect (ANBS).’ Despite the fact that it was found in September 1869, there appears to be no contemporary publication of this record. Dr A. N. B. Simpson very kindly Entomologist’s Gazette (2020) Vol. 71 227 examined the specimen again for us. It bears a small round data label with the following information: ‘bred/one Sept/69 from birch/Boughton/Worcester’. He also provided us with images, confirming the view expressed in the 2003 publication that it is Lyonetia clerkella. The next and final nineteenth century record was in 1893, when larvae were found by W. H. B. Fletcher at Worthing, Sussex. There are two published versions of this discovery. The first was by Tutt (1898: 199) who states: ‘Lyonetia padifoliella turned up at Worthing in some plenty in 1893, feeding on [Malus sp.], Cotoneaster affinis and Prunus japonica (sinensis) in August– September. On apple, the larvae patronised the topmost of the of the year. The lepidopterist, therefore should not summer-prune his apple-bushes (Fletcher).’ This record is repeated, but with two significant differences, by Fletcher (1905: 208–209) who, under L. prunifoliella, states that ‘In September, 1893, the larvae of this species were abundant in leaves of apple in the Infirmary garden, Worthing, and a few were found feeding also in those of Prunus sinensis in another garden in the same town (Fletcher)’. It is clear that Tutt’s note was based on Fletcher’s observations and so we do not know why Tutt cites Cotoneaster affinis Lindl. as a foodplant and gives the larval period as August–September since Fletcher does not mention this foodplant and gives only September for the larval period. We therefore treat Tutt’s inclusion of this as an error. We have traced over 40 specimens reared by Fletcher. All are L. prunifoliella f. albella. Details are given in the next section. There were no records in the twentieth century until 27 August 1974 when Emmet (1975a; 1975b) found a vacated leaf-mine on Sorbus aucuparia near Anstey, Leicestershire. Emmet exhibited this leaf-mine on two recorded occasions. The first was on 10 October 1974 at an ordinary meeting of The British Entomological and Natural History Society. The report of his exhibit (Emmet, 1975a) states: ‘A leaf-mine, believed to be that of Lyonetia prunifoliella ...’ He recognised the potential significance of the record as he comments that: ‘The species does not appear to have been recorded from Britain since the end of the nineteenth century or, possibly, the beginning of the twentieth. Meyrick gives the foodplants as blackthorn [Prunus spinosa] and birch [Betula sp.], but Hering also gives a wide range of , including Sorbus.’ The second occasion was about three weeks later on 2 November 1974 at the Annual Exhibition of the same Society. The report (Emmet, 1975b) states: ‘Lyonetia prunifoliella Hübn., a mine, evidently of this species ...’. A. M. Emmet was Britain’s foremost leaf-mining expert. We have examined his leaf-mine herbarium, which is in the Natural History Museum, London (NHMUK). This is arranged in alphabetical order under the scientific names of . Each leaf-mine is in a separate transparent envelope bearing a manuscript label in Emmet’s distinctive handwriting giving the scientific names of both the species of Lepidoptera and foodplant as well as other data. In the section for ‘Sorbus aucuparia’ is an envelope containing a short stem with one mine. The following determination is in Emmet’s handwriting: ‘Stigmella sorbi (Stainton)/Sorbus aucuparia/Anstey, Leics. 27.viii.74’ We examined the underside of the leaf which showed a typical nepticulid eggshell. Thus it is clear that after the publications indicating that he had found 228 Entomologist’s Gazette (2020) Vol. 71

Photo: R. J. D. I. Voith Fig. 5. Lyonetia padifoliella (Hübner, [1813]), white form. Norway: On, Nord-Fron, Vinstra, Stordalsberget ex Cotoneaster scandinavicus B. Hylmö 6.vi.2017.

Photo: R. J. D. I. Voith Fig. 6. Lyonetia padifoliella (Hübner, [1813]), dark form. Norway: On, Nord-Fron, Vinstra, Stordalsberget ex Cotoneaster scandinavicus B. Hylmö 6.vi.2017. Entomologist’s Gazette (2020) Vol. 71 229

Photo: L. Aarvik Fig. 7. Lyonetia padifoliella (Hübner, [1813]), dark form. Norway: On, Nord-Fron, Vinstra, Stordalsberget ex Cotoneaster scandinavicus B. Hylmö 20.vii.1983.

Photo: L. Aarvik Fig. 8. Lyonetia padifoliella (Hübner, [1813]), dark form. Norway: On, Nord-Fron, Vinstra, Stordalsberget ex Cotoneaster scandinavicus B. Hylmö 20.vii.1983. 230 Entomologist’s Gazette (2020) Vol. 71 a mine of L. prunifoliella, Emmet redetermined this as Stigmella sorbi. This must have been before publication in 1985 of MBGBI. Emmet (1985: 223) wrote the account of the Lyonetia, as well as that of the two British species, L. prunifoliella and L. clerkella, and states that these two species have been recorded in the British Isles although one ‘has not been observed for over 50 years.’ Therefore, up to 1985 Lyonetia prunifoliella had only been recorded three times in England, all in the nineteenth century. It was only found once in the larval stage, in 1893 at two gardens in Worthing, Sussex.

Consideration of the account of Lyonetia prunifoliella in The Moths and Butterflies of Great Britain and Ireland 2 In the account of this species Emmet (1985: 224) places Lyonetia padifoliella as a junior synonym of L. prunifoliella. South (1884: 39) was apparently the first to use the specific epithet prunifoliella in the British literature, placing the names padifoliella and albella as varieties of that species. Subsequently two British checklists (Kloet & Hincks, 1945: 138; 1972: 8) also treated Lyonetia padifoliella as a junior synonym of L. prunifoliella and, probably, Emmet was following these. Because Emmet lists L. padifoliella as a junior synonym of L. prunifoliella some of his account may relate to L. padifoliella, not least because of the lack of contemporaneous material that he could consult. Two adults are illustrated as figures 27 and 28 on plate 9 in that work. The illustrations of both show specimens with a considerable amount of brown on the forewing. The form of L. prunifoliella, having a mainly white forewing is not illustrated, nor is it described in the text. Figure 27 is almost certainly of L. padifoliella, whereas figure 28 is definitely of L. prunifoliella f. albella. We have not been able to trace the specimen used for figure 27 but the specimen from which the illustration was made for figure 28 is in the NHMUK, which we have examined. It bears three labels, in descending order: 1. ‘W.B.F./Worthing,/bred Oct.|93’ 2. ‘Ex Banks [sic] Coll.’ 3. A green rectangular label which has been cut so that not all the words are complete but which reads ‘Brian Hargreaves/Moths and Butterfli[es]/Britain and Ireland’. Brian Hargreaves was the illustrator of the colour plates in MBGBI. The reason that there is neither a description nor an illustration of the white form is presumably because none of the prior descriptions in the British literature accords with this form and the only figure of the adult in the British literature was that provided by Stainton (1848: fig. 30) which is of f. albella. Further, all of the extant English specimens known to us, over 40 and from no later than 1893, are of that form. These are in the NHMUK and the Collection, University Museum of Zoology, Cambridge (UMZC). In addition to the specimen used to illustrate figure 28 on plate 9, the NHMUK also has several other Fletcher specimens, all with data labels giving the month and year as October 1893, the locality as Worthing and indicating that they were bred, but no foodplant is given. There is also one Fletcher specimen with the month and year given as November 1893, the locality as Worthing but with no indication that it was bred. There are Entomologist’s Gazette (2020) Vol. 71 231

Photo: R. J. Heckford Fig. 9. Figure 30 in Stainton (1848) illustrating ‘padifoliella’.

Photo: Dr D. C. Lees Fig. 10. Lyonetia padifoliella (Hübner, [1813]), dark form. Switzerland: Bremgarten ex Betula sp. Frey collection, NHMUK.

36 specimens in the Fletcher Collection, in the UMZC. None has a data label but each has an oblong typed accession label reading ‘Ex coll. W. H./Ballett Fletcher/1941: 1’. In addition, one also has an oval manuscript label reading ‘S. Stevens/12.1857’. All except this specimen are standing under a manuscript label, presumably in Fletcher’s handwriting, reading ‘Worthing f1[?]. larvae/taken 13 & 27.9.[18]93’. The specimen with the Stevens label of 1857 stands below and to the left of this manuscript label. J. C. Stevens was the business name of a family of auctioneers that specialised in the sale of natural history specimens. Samuel Stevens was part of this business. He was also a keen collector of Lepidoptera, including . He lived from 1817–1899. His collection was sold in 232 Entomologist’s Gazette (2020) Vol. 71

2 parts totalling 876 lots, on 27–28 March 1900 and 23 April 1900, by his family business. If the specimen with the Stevens label is British then it is the earliest known extant one. If it is British then it seems likely that it must have been taken by Desvignes rather than Stevens otherwise surely Stainton or Stevens would have published the record. Desvignes lived from 1812–1868 and his collection was sold by J. C. Stevens on 30 June 1868. It is clear that Emmet saw the Fletcher material in Cambridge because he notes (1975a) that there is a bred series from Worthing, Sussex, in that collection. He almost certainly saw British material in the NHMUK because it is likely that he chose the specimen that is illustrated as figure 28. The account of the adult makes no mention of the antenna. This is because in the description of the genus Lyonetia the antenna is described as ‘slightly exceeding length of forewing’ (Emmet, 1985: 223). Although this is correct for L. clerkella, the antenna of L. prunifoliella significantly exceeds the length of the forewing as does that of L. padifoliella. Unfortunately in both figures 27 and 28 on plate 9 the antennae are erroneously shown to be about three-quarters the length of the forewing, as is the antenna of L. clerkella in figure 29 on that plate. Although it is possible that the specimens used for figures 27 and 29 may have had broken antennae, our examination of that used for figure 28 clearly has the right hand antenna distinctly longer than the forewing, although the left hand one does not because it is broken. As regards foodplants, Emmet (1985: 224) gives ‘various rosaceous and , including blackthorn (Prunus spinosa), Japanese (), hawthorn (Crataegus spp.), apple (Malus spp. including cultivars), Cotoneaster spp. and Sorbus spp.: also on birch (Betula spp.)’. In this list of foodplants given by Emmet ‘Cotoneaster spp.’ is included but not Cotoneaster affinis. We therefore assume that this entry is not based on Tutt (1898: 199) whose record of that plant we have already discounted for the reasons given in an earlier section. Therefore the inclusion of Cotoneaster spp. as well as Prunus spinosa, Crataegus spp. Sorbus spp. and Betula spp. must have been derived from observations made in mainland Europe. We now consider their possible source. The first mention of the larval foodplant in the British literature appears to have been by Stainton (1856: 58), who in turn published this on the basis of European information. He states, under Lyonetia padifoliella: ‘Professor Frey has lately sent me specimens of this insect (his Albella), of Prunifoliella, and of the Hübnerian Padifoliella, with the remark:—“I bred in August all these forms from one kind of sloe [Prunus spinosa] mine, with similar larvae, and am disposed to unite them all as one species. The mine is broad, not narrow like that of Clerkella. Albella is very scarce, Padifoliella the commonest. The pupa is suspended as in Clerkella.” After groping so long in the dark about this species, such a glare of light is rather dazzling!’ In the same year, Frey (1856: 311–312) gave an account under the name of L. prunifoliella in which he listed ‘L. Albella’ and ‘L. Padifoliella’ as varieties. Against L. Albella’ he gives ‘Padifoliella Sta. and ‘Acerifoliella Curt.’ In the text he gives the larval period as from the end of July to mid-August and cites the larval foodplant as Prunus spinosa. In a footnote, however, he comments that Bremi found it mining leaves of birch (Betula sp.) and that he, Frey, reared the padifoliella variety from birch provided by Bremi. Entomologist’s Gazette (2020) Vol. 71 233

The following year Frey (1857), again under Lyonetia prunifoliella, states that ‘Mines similar to those made by this insect in sloe leaves occur also on birch, rather sparingly. I have bred two specimens of the perfect insect.’ Stainton ([1857]: 112) cites Frey’s note, but under the name of Lyonetia padifoliella. We have examined two specimens from the Frey collection in the NHMUK. Each has a typed accession label and a manuscript label. The manuscript label of one reads ‘L. prunifoliella./var. albella./Eversm./Zürich.’, the other reads ‘Brem garten./el. von Betula.’. Both are clearly the dark form of L. padifoliella. The two localities given, Zürich and ‘Brem garten’, or rather ‘Bremgarten’, are both in Switzerland. Neither label has a captor’s name. Nevertheless, Frey (1880: 417), in a publication on the Lepidoptera of Switzerland, states that he recorded L. prunifoliella at Zürich and Boll found it at Bremgarten. He also observes that he noted both var. padifoliella and var. albella at Zürich. On this basis it may be that the specimen in Frey’s collection reared from Betula (Fig. 10) resulted from a larva collected by Boll. Meyrick (1895: 757; [1928]: 811) states ‘in blotches in leaves of blackthorn and birch’, and in both publications he gives ‘Northampton (Whittlebury Forest), local’ as the only locality. It would seem that he derived the foodplants from Frey’s publications. Hering (1957), in his account of European leaf-miners, only cites Lyonetia prunifoliella, and so, presumably, considered L. padifoliella to be a junior synonym. As they are now recognised as separate taxa this means that not all of the foodplants he lists for L. prunifoliella may be those of that species. Hering gives only generic names and these are: Betula (1957: 179–180, fig. 132), Chaenomeles (1957: 275–276), Cotoneaster (1957: 338), Crataegus (1957: 347–348), Cydonia (1957: 366), Malus (1957: 660, fig. 409), Mespilus (1957: 689), Prunus (1957: 837), Pyrus (1957: 851, fig. 409) and Sorbus (1957: 1010). Although he cites fig. 409 as being a figure of the mine on Pyrus, he also gives the same number for the figure of the mine on Malus. In fact fig. 409 shows the mine on Malus only. There is no illustration of the mine on Pyrus. The inclusion by Emmet (1985: 224) of ‘Japanese quince (Chaenomeles japonica)’ may have been based either on Tutt (loc. cit.) who gives ‘Prunus japonica (sinensis)’, Fletcher (1905: 208–209) who gives ‘Prunus sinensis’ or Hering (1957: 275–276) who gives ‘Chaenomeles’. Our attempt at determining both the source of the inclusion of this name and its current nomenclature has proved difficult and we accept that our view given in this section may not be definitive. First, a question arises as to whether Prunus japonica (sinensis) and Prunus sinensis are conspecific. According to World Flora Online (2020) Prunus japonica Thunb. is an accepted name. Prunus japonica (sinensis) is not listed, but ‘Prunus sinensis’ is listed twice, each name with a different author, and each with the comment that its status is ambiguous. Prunus japonica has various vernacular names including Dwarf flowering cherry and Oriental bush cherry. Chaenomeles japonica (Thunb.) Lindl. ex Spach is a different species, with Pyrus japonica Thunb. and Cydonia japonica (Thunb.) Pers. given as junior synonyms. Chaenomeles japonica has various vernacular names including Japanese Quince. We are extremely grateful to A. J. Halstead (in litt.) for the following information. Prunus and Chaenomeles species have very different . Those of Prunus have a single seed within the whereas those of Chaenomeles contain a number of seeds. Prunus japonica is apparently no longer commercially available 234 Entomologist’s Gazette (2020) Vol. 71

Photo: Miss S. D. Beavan & R. J. Heckford Fig. 11. Lyonetia prunifoliella (Hübner, 1796). England: Devon, Wembury, 7.x.2017, penultimate instar 3 mm long photographed 8.x.2017.

Photo: Miss S. D. Beavan & R. J. Heckford Fig. 12. Lyonetia prunifoliella (Hübner, 1796). England: Devon, Wembury, larva in mine found 7.x.2017, photographed 7.x.2017. Entomologist’s Gazette (2020) Vol. 71 235

Photo: Miss S. D. Beavan & R. J. Heckford Fig. 13. Lyonetia prunifoliella (Hübner, 1796). England: Devon, Heybrook Bay, 4.x.2017, final instar 5 mm long photographed 8.x.2017.

Photo: Miss S. D. Beavan & R. J. Heckford Fig. 14. Lyonetia prunifoliella (Hübner, 1796). England: Devon, Exminster, three larvae in mine found 22.ix.2017, photographed 24.ix.2017. 236 Entomologist’s Gazette (2020) Vol. 71 in the United Kingdom, although it may have been more widely grown in the 1890s. It is difficult to be certain whether Tutt and Fletcher were referring to a Prunus species or to what is now Chaenomeles japonica but it seems to us more likely that they were referring to Prunus japonica. As far as we can trace, Cydonia japonica is the name that had long been used in British horticulture before the change of the generic name to Chaenomeles and therefore they are more likely to have used that name if that was the species on which mines had been found. It seems very unlikely that either Jacq. or Crataegus laevigata (Poir) DC, the only native species of Crataegus in the British Isles, are larval foodplants in Britain if indeed they are foodplants of Lyonetia prunifoliella as opposed to L. padifoliella. This is based on our observations of larval mines at seven localities in Devon up to and including the time of writing this paper in August 2020. We have found over 400 mines, all on Prunus spinosa. Although Crataegus monogyna was always growing with Prunus spinosa in those localities we have never found mines on that. Therefore it appears that Emmet (1985: 224) was relying, at least in part, on the foodplants cited by Hering. Nevertheless, we do not know why Emmet did not include all the genera of plants given by Hering. We are not, however, implying that those not cited by Emmet are foodplants of Lyonetia prunifoliella. Emmet (loc. cit.) describes the larva as pale green and gives the larval period as July to August. He probably followed Meyrick (1895: 757; [1928]: 811) for this information. Emmet states that the pupa is undescribed. The species is given as univoltine with the adult emerging in late September, overwintering and on the wing again in the spring until May. Meyrick (1895: 757; [1928]: 811) gives ‘Larva pale green’ with the period for the adult as September and October. It is probable that Meyrick based his description on the detailed account by Frey (1856: 311–312), who, as already mentioned, treats Lyonetia padifoliella as a variety of L. prunifoliella. Frey states, in translation: ‘The larva is pale sea green, in slim, thin form similar to that of the following species [Lyonetia clerkella], with sharp and strongly separated from each other segments and a dark dorsal line. On the prothoracic plate are two black dots. The head brownish; the thoracic legs, close to each other, are black, whilst the prolegs have the greenish body color. The pupation is just like in the following species, L. clerkella.’ Frey does not specifically record that the description was made from a larva feeding on Prunus spinosa, but as this appears in his text as the foodplant we assume that the description was made from a larva feeding on that species. There is a slight caution, however, because as stated on page 232 in a footnote he also refers to Bremi finding the larva on Betula sp. As regards its English distribution, Emmet (loc. cit.) gives: Boughton, Worcestershire; Whittlebury Forest, Northamptonshire; near Stony Stratford, Buckinghamshire; Worthing, Sussex. References are provided for each record, which we have considered in an earlier section. Surprisingly, he comments that the species ‘was fairly common in the late nineteenth century’ at Worthing, citing Fletcher (1905) who does not state this. He records it only in September 1893, in two gardens in Worthing. Entomologist’s Gazette (2020) Vol. 71 237

English records of Lyonetia prunifoliella after 1985 and up to 2017 There were no confirmed records of L. prunifoliella in the twentieth century either before or after 1985. Thus it was a considerable surprise when after an absence of about 114 years, since 1893, the species was recorded on two occasions in 2007. Adults were found at Thurlbear, Somerset, on 17 August and at Puddletown, Dorset, on 13 October (Langmaid & Young, 2009: 21). The Somerset specimen is in the NHMUK which we have examined and it is the white form. Thereafter it has been recorded on several occasions. One was noted in a light-trap on the Lizard, Cornwall, when it was inspected on 2 October 2009 (Kitchener, 2010: 73–74). Two photographs of the live moth accompany the account and the individual is clearly the white form of L. prunifoliella. One was recorded at light at Elms Farm near Icklesham, Sussex, on 1 August 2013 (Pratt, 2015: 44). Langmaid & Young (2014: 208), presumably referring to the same record, give the date as 5 August and locality as Icklesham but do not mention Elms Farm. Then one white form came to light at Milton Common, Portsmouth, Hampshire on 7 August 2014. This was shown at the 2014 Annual Exhibition of the British Journal of Entomology and Natural History Society, where it was photographed (Thirlwell, 2015). Another of the same form was recorded at light in Sussex on 10 August 2014 in Old Wickhurst Lane in the Broadbridge Heath part of Horsham (Pratt, loc. cit.). On a date not given, a tenanted mine was found on Prunus spinosa, and moth reared, at Aylesham, Kent (Langmaid & Young, 2016: 291). The adult was recorded at Barnet, Middlesex on 18 July 2016 (Terry, 2018). One was ‘found sitting on a piece of Prunus in a back ’ at Catfield, Norfolk on 1 September 2017 (Wheeler, 2020). Three photographs of the live moth accompany that note which show that it is the white form. Then over 340 mines were found by the authors between 19 September and 7 October 2017 at various localities in Devon. The authors contacted M. S. Parsons about this which led to a visit by him on 5 October that year to Cogden Beach, Dorset where hundreds of mines were found, all on Prunus spinosa, with ca 5–10% being tenanted (Parsons, 2018). One was recorded at Lydd, Kent on 31 October 2017 (Clancy, 2018) and on the same date another was recorded by M. Opie at Horsea Island, Portsmouth, Hampshire (Dickson, 2018). These may well not be the only records of the species in England since 2007 and up to the end of 2017, but they are the only ones that we have traced.

Discovery of over 340 mines of Lyonetia prunifoliella on Prunus spinosa in Devon in 2017 Prior to 19 September 2017 Lyonetia prunifoliella had never been recorded in Devon. That day we visited the Exeter Canal area at Exminster Marshes, in south Devon to make records, mainly of leaf-mines. The first Prunus spinosa bush that we looked at had mines that neither of us had seen before, but were clearly those of Lyonetia prunifoliella. We found nearly 30 mines of which seven were tenanted. Encouraged by this, during the next couple of weeks we visited several other places, all in south Devon and all either on the coast or a few kilometres inland. The result was that we saw over 340 mines, with over 40 tenanted, at 10 sites up 238 Entomologist’s Gazette (2020) Vol. 71

Photo: Miss S. D. Beavan Fig. 15. Lyonetia prunifoliella (Hübner, 1796). England: Devon, Exminster Marshes, 19.ix.2017, larval mines.

Photo: Miss S. D. Beavan & R. J. Heckford Fig. 16. Lyonetia prunifoliella (Hübner, 1796). England: Devon, Exminster, 22.ix.2017, larval mines. Entomologist’s Gazette (2020) Vol. 71 239

Photo: Miss S. D. Beavan & R. J. Heckford Fig. 17. Lyonetia prunifoliella (Hübner, 1796). England: Devon, Heybrook Bay, larval mines 2.x.2018, photographed 4.x.2018 showing extruded frass.

Photo: Miss S. D. Beavan & R. J. Heckford Fig. 18. Lyonetia prunifoliella (Hübner, 1796). England: Devon, Exminster, larva found 22.ix.2017, pupa photographed 29.ix.2017. 240 Entomologist’s Gazette (2020) Vol. 71

to and including 7 October. Going from west to east the sites were at Jennycliff, Bovisand, Heybrook Bay, Heybrook, Wembury, Exminster and Exminster Marshes. We failed to find mines at Dawlish Warren and Budleigh Salterton. All the mines were on Prunus spinosa and towards the tips of stems on leaves that were slightly paler green than most of the leaves and looked ‘newer’. Most were on tips that were between about 1½ and 2½ m high above ground level. This reflects Fletcher’s observations, recorded by Tutt (1898: 199), that ‘On apple, the larvae patronised the topmost leaves of the shoots of the year. The lepidopterist, therefore, should not summer-prune his apple-bushes’. Moths emerged between 1 and 30 October 2017 and were the white form, f. albella and the intermediate form. In subsequent years we have found mines towards the tips of plants of Prunus spinosa that were no more than a metre above the ground.

Description of the early stages based on observations in Devon, England, in 2017 Ovum. Undescribed. The start of the mine is often adjacent to a very small hole in the leaf which may have been created when the larva emerged from the ovum. Not usually laid near the edge of the leaf. Larva. We are unclear about the number of instars. We did not observe any larva while in its linear mine. The descriptions below are based on larvae in blotch mines. Penultimate instar (Fig. 11). 3 mm long. Head, labrum and clypeus pale reddish brown, otherwise greyish black, adfrontal sutures slightly darker; prothoracic plate undivided, small, pale greyish black; thoracic and abdominal segments deeply incised; body very pale greenish white, pinacula inconspicuous; setae translucent, colourless; peritremes of spiracles un- described; anal plate small, pale greyish black; thoracic legs slightly translucent, pale greyish black, ventral and anal prolegs undescribed. The larva appears whiter when in the mine (Fig. 12). Final instar (Fig. 13). 5 mm long. Head, labrum and clypeus pale reddish brown, otherwise translucent showing body colour, adfrontal sutures very pale brown, stemmata black; prothoracic plate undivided, translucent slightly shiny showing body colour; thoracic and abdominal segments deeply incised; body pale green, often showing body contents darker green, a pair of elongate oval translucent slightly shiny pinacula on thoracic segments 2 and 3, otherwise pinacula inconspicuous; setae translucent, colourless except those on abdominal segments 9 and 10 which are pale greyish black; peritremes of spiracles inconspicuous, very pale yellowish brown; anal plate comparatively large, translucent very pale yellowish, very pale greenish or very pale brownish; thoracic legs undescribed, ventral and anal prolegs concolorous with body, crochets undescribed. The larva appears darker green when in the mine (Fig. 14). Mine (Figs 15, 16, 17). Initially a long, linear, often sinuous, mine almost filled with interrupted black frass; sometimes the mine follows veins, sometimes crossing these and the midrib, often commencing from a reddish brown area adjacent to a very small oval hole in the leaf; then abruptly widening into a comparatively large pale greenish blotch in which some of the black frass is scattered, but most is extruded in long thread-like strings and hangs down through one or more holes in the lower epidermis. Hering (1951: 36) records that changing mines is observed in this species, especially when the larva is changing from linear-mining to blotch-mining with the result that the initial mine is very often separated from the subsequent blotch, and the two are not infrequently in different leaves. We did not observe linear mines that did not lead to blotches, but we did observe blotches without linear mines. Sometimes up to three larvae were found in one blotch mine suggesting that either that ova were laid close together or that sometimes larvae somehow ‘congregate’. In old mines the frass often becomes reddish brown and the blotch mine becomes pale yellowish brown or pale brown. Pupa (Figs 18, 19, 20). 4 mm long. Pale green to slightly darker green becoming very pale yellowish white posteriorly, suffused greyish black on pair of distinct rather pointed cephalic Entomologist’s Gazette (2020) Vol. 71 241 horns and at distal ends of all legs, antennae exceeding abdomen by about 1.5 mm, green annulated pale greenish grey from distal ends of hindlegs becoming darker and eventually greyish black beyond abdomen, gonads yellowish. Pupa eventually showing colour and markings of moth. In a flimsy whitish cocoon, suspended at both ends by several long silken threads that were usually attached to parts of the containers within which the larvae had been kept, but sometimes the cocoons were spun below the leaves. Exuviae translucent, colourless except for the areas suffused greyish black in the pupa which are pale greyish black, not extruded from cocoon on emergence. When the moths emerged, they usually initially rested amongst silken strands near the exuviae (Fig. 21). The pupa of Lyonetia clerkella is very similar (Fig. 22), both in the colour of the pupa as well as the suffused greyish black markings at the distal ends of all legs, and with the antennae pale grey from the distal ends of the hindlegs becoming darker and eventually greyish black beyond abdomen and the gonads yellowish, but the cephalic horns are noticeably smaller and more blunt and are not suffused greyish black and the antennae only exceed the abdomen a little way (not measured). The pupa is in a similar cocoon and has similar strands attaching it to the substrate.

Consideration of larval foodplants of Lyonetia prunifoliella and L. padifoliella Although Hübner recognised Lyonetia prunifoliella and L. padifoliella as different species they were synonomized by Wocke (1871: 333), and followed by South (1884: 39) in the British literature. Thereafter L. padifoliella continued to be treated as a junior synonym in both British and European literature. For example, L. padifoliella is not mentioned by Baraniak (1996) in his account of the Lyonetiidae in the distributional checklist of the Lepidoptera of Europe. In 2011 Bengtsson & Johansson (2011: 421–423) showed that it was a good species. Therefore prior to 2011 not all the foodplants attributed to Lyonetia prunifoliella might be of that species. Bengtsson & Johansson (loc. cit.) cite only Prunus spinosa for L. prunifoliella and Cotoneaster scandinavicus for L. padifoliella, giving the Nordic distribution for the latter as only southern Norway. Later, however, although Aarvik et al. (2017: 161) still record only Cotoneaster scandinavicus in Norway for L. padifoliella, they state that recent studies by Dr L. Kaila have shown that the species also occurs in Finland where it has been bred from ‘Betula, Malus, Sorbus, Cotoneaster and Aronia. (M. Mutanen, L. Sippola, E. Saarela, pers. comm.). DNA barcoding of Norwegian and Finnish specimens has confirmed that they are conspecific.’ They do not cite larval foodplants for L. prunifoliella, but this is not surprising as their publication is a checklist, with relevant notes, and not a field guide. Dr Kaila tells us (in litt.) that both Prunus spinosa and Prunus cerasus are among the verified host- plants of L. prunifoliella but that in Japan and Korea L. prunifoliella ssp. malinella Matsumura, 1907, feeds on Malus. This taxon shares the barcode and morphology of the European Prunus feeder. Subsequent to those publications, Agassiz & Langmaid (2018: 99) give ‘Prunus spinosa, Chaenomeles japonica, Betula spp., Malus or Cotoneaster spp.’ as larval foodplants of L. prunifoliella in England. None of these names is preceded by an asterisk (*). If any had been it would mean that the information was derived from a mainland European source. Those are the same foodplants given by Emmet (1988: 51) where the same asterisk convention is employed. 242 Entomologist’s Gazette (2020) Vol. 71

Photo: Miss S. D. Beavan & R. J. Heckford Fig. 19. Lyonetia prunifoliella (Hübner, 1796). England: Devon, Heybrook Bay, larva found 4.x.2017, pupa photographed 10.x.2017, ventral view.

Photo: Miss S. D. Beavan & R. J. Heckford Fig. 20. Lyonetia prunifoliella (Hübner, 1796). Same pupa as in fig. 19, lateral view. Entomologist’s Gazette (2020) Vol. 71 243

Photo: Miss S. D. Beavan & R. J. Heckford Fig. 21. Lyonetia prunifoliella (Hübner, 1796). England: Devon, Exminster, larva found 22.ix.2017, moth emerged freshly emerged 5.x.2017, with exuviae above it.

Photo: Miss S. D. Beavan & R. J. Heckford Fig. 22. Lyonetia clerkella (Linnaeus, 1758). England: Devon, Heybrook Bay, pupa, photographed 10.x.2017. 244 Entomologist’s Gazette (2020) Vol. 71

For the reasons given earlier in this paper, we consider that the only confirmed larval foodplants in England are Prunus spinosa, Prunus japonica and Malus sp., and so far only found once on each of the last two. It is probable that it may also be found on other Prunus species.

Consideration of whether any English records of Lyonetia prunifoliella are of L. padifoliella It is clear from figure 30 in Stainton’s 1848 paper adding ‘padifoliella’ to the British list that in fact the species was Lyonetia prunifoliella f. albella. As previously discussed, we have not traced any information about the record from near Stony Stratford, Buckinghamshire. The specimens reared in 1893 from Worthing, Sussex are also L. prunifoliella f. albella. As regards records published this century that are not accompanied by a photograph we cannot definitely determine the issue. In view of the considerable number found in 2017, however, and an increasing influx of the species since then on balance it seems unlikely that any are of L. padifoliella.

Summary and discussion In the light of subsequent information as well as reconsideration of the information available at the time in 1985 we suggest that parts of the account of Lyonetia prunifoliella by Emmet (1985: 224, pl. 9 figs 27, 28) in MBGBI should be read in conjunction with the following observations. The description of the adult does not include a description of the form having mainly white forewings, and as a consequence plate 9 does not include this. Neither the description of the adult, nor the generic description of the Lyonetiidae, mentions the fact that the antenna significantly exceeds the length of the forewing, and the illustration on plate 9, figure 28 wrongly shows the antenna to be significantly shorter than the forewing. Figure 27 on plate 9, now known to be that of L. padifoliella, also shows the antenna to be significantly shorter than the forewing, whereas it also exceeds that length. Although the larval period is given as July to August this does not agree with the only English record of larvae known at the time of that publication. Larvae were found in ‘August–September’ or simply ‘September’ in 1893. Since 1985 larvae have been found between at least July to October, on the basis of various online reports and as yet unpublished information. As regards foodplants, we refer to the last paragraph in the penultimate section above. Emmet (loc. cit.) comments that the species is stated to be univoltine with the adult emerging in late September, overwintering and on the wing again in the spring until May. At the date of that publication there was no evidence in England that the species overwintered or that it continued as an adult until late May. Up to 1985, the only record of the adult where a month was given was in September, in Whittlebury Forest, Northamptonshire, taken by Desvignes. Since 1985 adults have been recorded between at least July and October, with one found on 6 January 2019 in Dorset (M. S. Parsons, in litt.). It is beyond the scope of this paper to conside in any detail the mainland European distribution of both Lyonetia prunifoliella and L. padifoliella. This would Entomologist’s Gazette (2020) Vol. 71 245 require, amongst other things, an assessment of the accuracy of historic records given under the name of either species. On the limited information obtained during the preparation of this paper it would seem that L. prunifoliella is the more widespread. The only confirmed records of L. padifoliella that we have traced are from Norway, Finland and Switzerland, but we suspect that it occurs elsewhere. More investigations are needed to assess the distribution and larval foodplants of both species. Finally there is the question as to whether the species merited pRDB1 status for the period 1980–2011 and whether it should still be regarded as a very rare resident. Parsons (2003: 64; 2010: 64) considered that it was an extinct resident, although noting the 2007 records in his later paper but, in our view correctly, commenting that these may have been primary immigrants. The sporadic nature of the records since the species was added to the British list coupled with the fact that larval mines in England have been almost exclusively found on the widespread and generally common Prunus spinosa, rather suggests that it is an immigrant that occasionally temporarily colonises to the extent of producing at least one generation or more. Prunus spinosa is frequently searched for the larvae of various species of Microlepidoptera. As a result, it seems very unlikely that Lyonetia prunifoliella was an overlooked resident before its rediscovery in 2007. The fact that leaf-mines have been increasingly found from 2017 does not necessarily mean that there are now resident populations in all those places. Our own experience in Devon shows that at some sites where larvae have been found in one year there are no mines the following year. We have, however, found mines at one locality from 2017 to 2020 which does suggest a temporary population over several years. It will be interesting to see if the species becomes permanently established there or in other places in England. In any event it is clear from the increasing number of records from various counties that the species can no longer be regarded as having pRDB1 status.

Acknowledgements We thank Mr M. S. Parsons (Walditch), Dr K. Sattler (NHMUK), Dr A. N. B. Simpson (Tenbury Wells) and Mr I. R. Thirlwell (Southsea) for their help in various ways. We are especially indebted to Sir A. C. Galsworthy (NHMUK) for providing us with translations of papers written in Chinese, which although not cited have proved extremely helpful, Mr A. J. Halstead (Woking) for his advice on some of the Prunus/Chaenomeles problems and Dr L. Kaila (Finnish Museum of Natural History, Helsinki) for providing us with information about certain host-plants as well as barcode and morphology of the species in Japan and Korea. We are extremely grateful to the following for photographing material for us: Dr D. C. Lees (NHMUK) for the image of the specimen from the Frey collection; Mr E. C. Turner (UMZC) for several images from the W. H. B. Fletcher collection; Mr L. Aarvik (Natural History Museum, University of Oslo) and Mr R. J. D. I. Voith (Lillehammer) for images of reared specimens of Lyonetia padifoliella from Norway from their collections and doing so at fairly short notice. We acknowledge the copyright of the latter two in respect of their photographs and the photograph from the Frey collection as belonging to the Trustees of the NHMUK. We thank the Trustees of the NHMUK and Dr Lees for allowing our use of that photograph. 246 Entomologist’s Gazette (2020) Vol. 71

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