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Entomologist’s Rec. J. Var. 131 (2019) 105 OBSERVATIONS ON THE LARVA AND FOODPLANTS OF RELIQUANA (HÜBNER, [1825]) (LEP.: , OLETHREUTINAE)

¹ S. D. B EAVAN AND ² R. J. H ECKFORD ¹ The Hayes, Zeal Monachorum, Devon EX17 6DF ² Department of Life Sciences, Division of , Natural History Museum, Cromwell Road, London SW7 5BD (Correspondence address: 67 Newnham Road, Plympton, Plymouth, Devon PL7 4AW)

Abstract This paper considers accounts of the larva and larval foodplants of Lobesia reliquana (Hübner, [1825]) in British and mainland European literature as a result of finding larvae in withered brown leaves of Hornbeam Carpinus betulus L. and Quercus sp. in Devon, England. Certain cited foodplants are considered to be erroneous or requiring confirmation. Only a few larval descriptions are considered to be reliable. Key words : , Tortricidae, Lobesia reliquana , larva, foodplants, Carpinus betulus , Quercus sp., dead leaves.

Introduction Lobesia reliquana (Hübner, [1825]) is a comparatively widespread species in Great Britain although becoming more local or absent in northern England and Scotland. In view of this, it is perhaps rather surprising that there appear to be only three published accounts of the larva being found in the wild in Great Britain. The first was in 1887 by J. H. Wood (1888), who beat two larvae from Blackthorn Prunus spinosa L. By 1895 Wood (1895: 159) had recorded it from Betula sp. and then by 1992 (Robbins, [1992]: 166) it had been found on Wild Cherry Prunus avium (L.) L. Between 2009 and 2013 we collected several larvae that all appeared to be the same amongst withered brown leaves of Hornbeam Carpinus betulus L. and oak Quercus sp. that were attached to twigs that had been had been caught amongst branches in the understorey at two woods in Devon. We succeeded in rearing one in 2010 and another in 2014. Both proved to be Lobesia reliquana . Further details are given later in this paper. We examine a number of published records of the larval foodplants and suggest that eight of them, several first given in mainland European literature and then cited in British literature, are either erroneous or unconfirmed. We review the few published British larval records, as well as some from mainland Europe, none of which suggests that the larva feeds on dead leaves. Several of the earlier publications give the specific name of the species as ‘ permixtana ’ but it is clear from the context that it is Lobesia reliquana that is being referred to and not permixtana ([Denis & Schiffermüller], 1775) which is also in the Tortricidae. We provide a photograph of one of the larvae that we Ent Rec 131(3).qxp_Layout 1 25/05/2019 10:19 Page 106

106 Entomologist’s Rec. J. Var. 131 (2019) found because although it agrees with the original British description given by Wood (1888) it differs in several significant respects from that provided by Bradley, Tremewan & Smith (1979: 68) and Hancock & Bland (2014: 22).

Larval foodplants given in British and mainland European literature The following lists larval foodplants given in both British and mainland European literature in the order in which they were published. We follow Stace (2019) for both the scientific and vernacular names of the foodplants given, except for names included in quotations and also for the vernacular name of Juniperus oxycedrus L. because this does not occur in the British Isles and so is not included by Stace. Common Juniper Juniperus communis L. This is cited by Hartmann (1868; 1870: 36; 1880: 44). In his first publication he records that he reared one specimen of ‘Lobesia Permixtana HS’ as well as other species from ‘die Knoten’ on Juniper communis . In his second publication he states, ‘an Juniperus communis in aufgetriebenen Knoten der Stämme und Zweige’ (‘on Juniperus communis in swollen knots on the stems and branches). In his third publication he simply gives ‘ Juniperus communis ’ without any further information. Stainton (1887) refers to Hartmann’s 1870 publication and describes the record of the larva occurring in the swollen knots on the stems and branches of Juniperus communis as ‘somewhat startling information’, observing that ‘surely some other species must here be meant.’ Walsingham (1887) refers to Stainton’s note and comments it was extremely probable that the larva must have been that of Grapholita opulentana Millière, 1875 (now Cydia interscindana (Möschler, 1866)). He states that although he is not aware that the larva of that species feeds on Juniperus communis , he had frequently bred it from swollen knots on Prickly Juniper Juniperus oxycedrus L. Joannis (1921) carried out an extensive review of the literature dealing with the larval foodplants of Lobesia reliquana , including Hartmann’s 1868 publication. Joannis translates Hartmann’s ‘Knoten’ as ‘galles’ and concludes that the species was not cecidogenic (gall-forming). Lhomme (1935: 389) also states that it is not cecidogenic, but adding that it has been obtained accidentally from a gall of Juniperus communis , citing ‘Hartmann’. We entirely agree that Hartmann was mistaken. In our view it is inconceivable that the larva of Lobesia reliquana , which undoubtedly feeds on leaves of at least 10 different species of deciduous trees, and in our experience also in spun withered leaves, also feeds in swollen knots/galls in stems and branches of Juniperus communis . Alkanet Anchusa officinalis L. Jourdheuille (1870: 127) gives this, stating simply ‘Sur Anchusa officinalis ’. Stainton (1887), however, points out that Rössler (1881: 247) assigns this foodplant to Lobesia artemisiana (Zeller, 1847). Goldenrod Solidago virgaurea L. Brischke (1876: 68) found one larva ‘in den Stengelspitzen der Solidago virgaurea’ (‘in the tips of the stem of Solidago virgaurea ’) on 21 August 1871, with the moth resulting on 11 April 1872. Stainton Ent Rec 131(3).qxp_Layout 1 25/05/2019 10:19 Page 107

Entomologist’s Rec. J. Var. 131 (2019) 107 (1887) remarks that although Brischke’s note was not published until nearly four years after the appearance of ‘ the first permixtana reported to have been bred’ he does not appear to have found any more larvae. Stainton’s use of italics for the passage cited suggests that he had doubts about some aspect of the record, especially as he comments that ‘The observation may be a good one, but it is desirable to have it confirmed by those who have opportunities of collecting in August amongst Solidago .’ Fenn (1887) refers to Stainton’s note and states that in his district in Kent, England, the moth is abundant and flies over the tops of oak ( Quercus sp.) bushes and small trees. He comments that he can hardly think that the larva feeds on Solidago virgaurea as the moth is very common in woods where that plant is absent and that in collecting the stems and seed-heads for rearing various species of Lepidoptera he had never heard of anyone breeding this species, which in his experience was confined to oak. Sheldon (1920) obtained ova from captive females of Lobesia reliquana . Only one larva resulted and this fed on spun leaves of Quercus sp., but in the final instar Sheldon introduced a leaf of Solidago virgaurea . He records that although this leaf was then spun to the only leaf of Quercus sp. in the container, it was not fed upon. As far as we are aware there is no other record of the species being bred from Solidago virgaurea . We consider that Brischke had simply found a larva that had descended from higher up in order to pupate. This assumes that Brischke’s indentification of the resulting adult was correct. Beech Fagus sp. We presume that the following refer to Fagus sylvatica L. even though only ‘ Fagus ’ is given. Sorhagen (1886: 104) gives the larva as occurring in August in the tips of the stems of Solidago virgaurea (Brischke) , Anchusa officinalis (Jourdheuille) and woody knots of Juniperus communis according to Hartmann, and also on deciduous trees, according to Sauber in September on Fagus . In his Introduction, Sorhagen (1886: viii –ix) lists a number of publications that he has consulted. The entry for Sauber is given as ‘Sauber, in den Hamburger Verh. 1871–74.’ The only relevant publication by Sauber that we can trace is in Verhandlungen des Vereins für Naturwissenschaftliche Unterhaltung zu Hamburg published in 1875 for the period 1871–1874. There Sauber (1875: 155) simply states: ‘Lobesia Gn. permixtana Hb. – S. – Ende Mai. Haake.’; no foodplant is mentioned. ‘Fagus ’ is subsequently cited by various authors, but we cannot trace a first-hand account of the larva being found on the leaves of this tree. Blackthorn Prunus spinosa L. This appears to be derived from Wood (1888) who beat two larvae from bushes of Prunus spinosa at the end of July 1887. In captivity the larvae were given only the leaves of Prunus spinosa ; there is no suggestion that these were withered. Benander (1965: 5) also gives ‘Prunus spinosa, larva 26/7; imago 16/5.’ We note that he does not provide a larval description whereas he does for most of his larval accounts. This was the second part of two papers that he published on the larvae of Swedish Microlepidoptera, all based on his personal observations of the previous 50 years. Downy Birch Betula pubescens Ehrh. Wood (1895: 159) records that ‘Birch ( Betula glutinosa ) [now Betula pubescens ] is a food-plant for this Tortrix , as well as Prunus Ent Rec 131(3).qxp_Layout 1 25/05/2019 10:19 Page 108

108 Entomologist’s Rec. J. Var. 131 (2019) spinosa . I have obtained it on several occasions from the plant. A suggestion thrown out that it may feed on oak is, therefore, likely enough some day to come true.’ He does not specifically state that he found larvae feeding on Betula sp., but this must be the implication because Wood was a careful observer. Further, a few years later in his account of the Lepidoptera of the county of Hereford he states (1908: 93) that ‘the larva feeds on sloe [ Prunus spinosa ] and birch [ Betula sp.], and probably on other plants’. Grey Alder Alnus incana (L.) Moench. Crombrugghe de Picquendaele (1906: 117) states that the larva is polyphagous and that he had taken it on Alnus incana on 8 August 1903, moth reared on 5 May 1904, and on ‘ bouleau’, Betula sp., on 10 September 1904 and this larva was described by Disqué (1904: 231). The determination of the larva found on Betula sp. was presumably based on the larva found on Alnus incana but no moth was reared because it is stated that the larva escaped. We set out in the next section the larval description that was given. This does not agree with undoubted larval descriptions of Lobesia reliquana . Elm Ulmus sp. Kennel (1916: 462–463) appears to be the first to cite Ulmus , but we cannot find any subsequent publication that mentions this. Oak Quercus sp. Sheldon (1920), after reviewing several publications, states that ‘There can be no doubt but that the imago has a strong penchant for oak [ Quercus sp.].’ On 4 June 1918 he caught several females resulting in ova being laid mainly on the upperside, but a few on the underside, of a leaf of Quercus sp. Just over 40 pages later in the same journal Thurnall (1920) records beating two larva in July ?1916 from ‘a shrubby oak’ in Brentwood, Essex. One moth was reared the following year. Hazel Corylus avellana L. Adkin (1931: 47) gives, ‘Larva in July and August on birch, hazel, etc.’ In the Preface to this publication about the of Eastbourne, Sussex, Adkin makes clear that he compiled his list from a number of sources and that W. G. Sheldon, an acknowledged expert on the Tortricidae and who elucidated the life- histories of a number of species in that family, is thanked for perusing the sections relating to the Pyralidae and Tortricidae. Although, as mentioned above, Sheldon reared larvae of Lobesia reliquana on leaves of Quercus sp. from ova laid by captive females, we cannot trace any published record by him of finding, or rearing, larvae on Corylus avellana . Nor have we traced any first-hand account of the larva being found on Corylus avellana by anyone else, and so we view this as a foodplant that requires confirmation. Alder Alnus glutinosa (L.) Gaertn. Lhomme (1935: 388–389) states that the larva is polyphagous on ‘ Prunus spinosa L., Betula alba L. [now Betula pendula Roth], Alnus glutinosa Gaertn., Fagus .’ We cannot find that Alnus glutinosa has been subsequently cited as a foodplant. Silver Birch Betula pendula Roth. As mentioned in the previous paragraph, Lhomme (1935: 388–389) gives this as a foodplant. Wild Cherry Prunus avium (L.) L. Robbins ([1992]: 116) gives ‘In spun leaves of Quercus and Prunus avium , the latter a new British host, (also on P. spinosa and Ent Rec 131(3).qxp_Layout 1 25/05/2019 10:19 Page 109

Entomologist’s Rec. J. Var. 131 (2019) 109 Betula ).’ In the Introduction to this publication Robbins comments on page 8 that as regards the foodplants cited, ‘Priority has been given to local observations, which are listed first, while any additional data from the literature are placed in parentheses.’ Hornbeam Carpinus betulus L. and Willow Salix sp. Razowski (2003: 23) lists these, as well as other larval foodplants given in this section. We have not been able to trace the source of either, but on 12 July 2009 we found three similar larvae, one amongst a withered brown leaf of Carpinus betulus and two amongst withered brown leaves of Quercus sp., at Budshead Wood, Devon and one Lobesia reliquana resulted from the larvae amongst Quercus sp. leaves. Bramble Rubus ulmifolius Schott . Corley et al . (2018: 563) record the species as new to Portugal on the basis of a moth reared on 23 April 2018 whose larva was collected on ‘ Rubus ’ on 17 September 2017 by J. Nunes. The latter has told us ( in litt .) that the Rubus species was almost certainly Rubus ulmifolius , the more common species of this genus in Portugal, and that the larva was feeding on young leaves in the understorey. He very kindly provided us with excellent unpublished images taken by him of the larva and also of a leaf of Rubus ulmifolius with part of the edge cut away and folded down onto the uppersurface of the leaf, where it was held by silk and within which the larva pupated. In addition to the foodplants cited in the literature mentioned above, Lepiforum (2019) provides two images of a larvae on Quercus robur L. and three images of a pupa resulting from a larva found on Betula pendula , one image of a pupa resulting from a larva found on Carpinus betulus together with an image of a leaf of that species, partly cut away and folded down onto the surface, being the pupation site, and one image of a similar cut-out and folded down leaf of Dogwood Cornus sanguinea L., also being a pupation site of a larva found feeding on this species. Several publications mention one or more of these foodplants but are apparently based on prior records as opposed to original observations. This can lead to the repetition of erroneous records and so we have not included them under each species of foodplant. However, we cite the following because they contain additional information. Bradley, Tremewan & Smith (1979: 68) comment on the larva ‘feeding chiefly on Quercus but also on Prunus spinosa and Betula , living in the terminal shoots or in spun leaves. On the Continent recorded also on Fagus , Solidago virgaurea and Anchusa officinalis (Hannemann, 1961: 199) and Juniperus .’ Falck & Karsholt (1998: 124) state that the main host-plant is Quercus , ‘but larvae … have also been recorded from Anchusa , Betula , Fagus , Juniperus , Prunus and Solidago ’, citing Bradley, Tremewan & Smith. Falck & Karsholt make the interesting comment that some of these host-plants may eventually be found to refer to Lobesia virulenta Bae & Komai, 1991, a species that was described from Japan and not recognised in Europe before 1997. Falck & Karsholt determined that the material of this species in Europe was sufficiently Ent Rec 131(3).qxp_Layout 1 25/05/2019 10:19 Page 110

110 Entomologist’s Rec. J. Var. 131 (2019) different to warrant subspecies status which they named Lobesia virulenta mieana . At the time of their publication this subspecies was known only from Denmark and Norway. Within a few years, however, it was additionally reported from Scandinavia, Poland and Slovakia (Razowski, 2003: 23). It is as yet unknown from the British Isles. Although, as far as we are aware, the larva of this subspecies has not been discovered, Falck & Karsholt (1998: 119, 121) state that the adults have been found in mixed deciduous forests with some being beaten from Bird Cherry Prunus padus L. Kubasik, Larsen & Chrzanowski (2002: 22) comment that Lobesia virulenta ‘most likely lives on ( Betula sp.), whereas L. reliquana lives mostly on ( Quercus sp.).’ Svensson (2006: 87–88) makes the interesting observation that ‘The life history of the larva is still not entirely investigated, but it can be surmised to live on Quercus in July.’ In The Moths and Butterflies of Great Britain and Ireland volume 5 (2) Hancock & Bland (2014: 22) record that the ovum is: ‘Laid in June on either surface of a leaf of oak ( Quercus spp.) (Sheldon, 1920a), or less frequently blackthorn ( Prunus spinosa ) or birch ( Betula spp.). On the Continent also recorded on beech ( Fagus sylvatica ), goldenrod (Solidago virgaurea ) and alkanet ( Anchusa officinalis ) (Hannemann, 1961), and juniper ( Juniperus communis ).’ The general format of The Moths and Butterflies of Great Britain and Ireland is to set out foodplant names in the section dealing with the ovum. This can sometimes give the misleading impression that the ovum has been observed to be laid on that plant, which in turn can give more credence to the plant cited as actually being a foodplant. The reference to ‘Sheldon, 1920a’ is to the same paper cited in this article as Sheldon (1920). The wording of this section by Hancock & Bland (2014: 22) relating to foodplants is almost exactly the same as the wording by Bradley, Tremewan & Smith (1979: 68). Therefore we presume that the foodplants given by Hancock & Bland are taken directly from Bradley, Tremewan & Smith and are not a separate independent assessment of whether they have been confirmed. Elliott et al. (2018: 245) give Quercus spp., Betula spp. and Prunus spinosa as larval foodplants, with the comment that the larva occurs ‘in terminal shoots or spun leaves. Overwinters full-fed from September in a cocoon spun in a dead leaf, in a spongy oak gall, or on the ground.’ The second sentence is based on otherwise unpublished observations of Dr J. R. Langmaid. He reared a few adults from galls of Biorhiza pallida L. (Hymenoptera, Cynipidae) collected on 28 August 1986 in Hampshire in the company of Lt. Col. D. H. Sterling and Dr P. H. Sterling. On 12 May 1984 Dr Langmaid reared a moth from leaf litter collected on 18 April that year at a different site Ent Rec 131(3).qxp_Layout 1 25/05/2019 10:19 Page 111

Entomologist’s Rec. J. Var. 131 (2019) 111 in Hampshire. Trees around the area where the leaf litter was collected were Aspen Populus tremula L. , Betula sp., Goat Willow Salix caprea L. and Pedunculate Oak Quercus robur L. Although Bradley, Tremewan & Smith (1979: 68), Razowski (2003: 23). Sterling & Parsons (2012: 304), Hancock & Bland (2014: 22) and Elliott et al. (2018: 245) all comment that the larva occurs ‘in terminal shoots’ we have not found any publication prior to Bradley, Tremewan & Smith ( loc.cit .) that suggests that the larva feeds in this way. It is unclear whether this comment was based on the observations of one or more of the authors of the 1979 publication or of one or more of the British Microlepidopterists who were consulted in the drafting of that publication.

Larval descriptions given in British and mainland European literature Brischke (1876: 68) appears to have been the first to describe the larva, if what he found was indeed the larva of Lobesia reliquana . He states, in translation: ‘On August 21st 1871 I found at Oliva in the tips of the stems of Solidago virgaurea a Tortrix larva, about 8 mm long, pale brown-green or brown-red, head and first segment shining brown, the latter with a pale central streak. Body set with individual hairs; glossy pinacula arranged on each segment on each side two, one diagonally behind the other, and two even more laterally, one standing above the other, arranged slightly differently on the 3rd segment. A nal shield shining brown. For pupation the larva goes into the earth and spins an elongate cocoon. The adult emerged on the 11th April 1872.’ As already noted, we consider that it is very unlikely that the larva was feeding on Solidago virgaurea . Brischke comments that the larva is pale brown-green or brown-red but as there was only one larva we do not understand the reason for giving alternative colours. Further, he makes no mention of the pinacula being white even though these are conspicuous on thoracic segments 2–3 although sometimes less conspicuous on abdominal segments 1–8. Taking all these points into consideration, we cannot be sure that the larva he found was that of Lobesia reliquana . Wood (1888) appears to have been the first to describe the larva in the British literature, and possibly the first to provide a larval description if Brischke’s larva was not that of Lobesia reliquana . He obtained two larvae by beating bushes of Prunus spinosa at the end of July 1887. In captivity the larvae were given only the leaves of Prunus spinosa ; there is no suggestion that these were withered. One moth emerged on 20 May the following year; the other died in the pupal stage. Wood describes the larva as follows: ‘The larva is cylindrical, of moderate proportions, and tapering moderately towards either extremity. The segmental divisions deeply cut; viewed from Ent Rec 131(3).qxp_Layout 1 25/05/2019 10:19 Page 112

112 Entomologist’s Rec. J. Var. 131 (2019) above, the profile of the segments is flat, not curved as is usually the case. Head small, shining, pale brown; thoracic plate also pale brown and shining, and with a narrow grey edging behind. Anal plate not noticeable. Colour a dark puce, with the spots [pinacula] white and small, but conspicuous. Legs black. Very active, jumping about like a Gelechia . The pupa is uniformly pale yellow, and not remarkable in shape.’ Disqué (1904: 231), under Lobesia permixtana , gives ‘5–6, Selten. Von Baron de Crombrugghe de Picquendaele in Brüssel ich Mitte Sept. eine R. von Betula. die mir aber leider entwischte. Sie war grünlich, der Rücken trüb braunrot. Kopf und Nackenschild hellbraun’ ( From Baron de Crombrugghe de Picquendaele in Brussels, in the middle of September a larva on Betula which unfortunately escaped. It was greenish. The back dull brownish red. Head and prothoracic plate light brown). As no moth was reared, because the larva escaped, it is not clear why the larva was con sidered to be that of Lobesia reliquana . Perhaps it was because Crombrugghe de Picquendaele (1906: 117) had found a larva on ‘bouleau’, Betula sp., on 10 September 1904 and the resulting moth was this species, and so he considered that this larva was the same. However, the description of the larva as greenish with the dorsum brownish red and with no mention of distinct pale pinacula does not agree with undoubted larval descriptions of this species. Barrett (1907: 248) provides a larval description based on Wood’s 1888 publication. Kennel (1910: 272) states that the larva is ‘dunkel purpurrot, mit kleinen weißen Wärzchen, Kopf und Nackenschild sind blaßbraun’ (‘dark purple-red, with small white pinacula, head and prothoracic plate are pale brown’). Although this is similar to the description given by Wood (1888) he does not cite Prunus spinosa as a foodplant. This suggests that the description may have been based on one that we have not traced because the foodplants he gives are the following, and presumably taken from the published literature cited in the previous section: the tips of the stem of Solidago virgaurea and Anchusa officinalis , also on deciduous trees, such as Betula , Fagus , also in woody knots on Juniperus . A few years later Kennel (1916: 462 –463) describes the larva as ‘grünlich, auf dem Rücken trüb braunrot, oder auch ganz dunkel braunrot, mit kleinen weißen Wärzchen, Kopf und Nackenschild sind hellbraun’ ( ‘greenish, on the back dull brown-red, or else completely dark brown-red, with small pinacula [colour unstated], head and prothoracic plate light brown’). I t is unclear why this description differs from that given by him six years earlier. It appears that the first part of the description may have been taken from Disqué ( loc. cit .), but it is unclear why the second part differs from his 1910 description which certainly applies to Lobesia reliquana . Ent Rec 131(3).qxp_Layout 1 25/05/2019 10:19 Page 113

Entomologist’s Rec. J. Var. 131 (2019) 113 The next British larval record that we can trace after Wood’s 1888 account is that given by Sheldon (1920). He states that on 4 June 1918 ova were laid by captive females mainly on the upperside, but a few on the underside, of a leaf of Quercus sp. (apparently the unstated number of females all laid ova on the one leaf). Sheldon gives a detailed account of the ovum, larva and pupa. Only one larva resulted from the ova. This was given three leaves of Quercus sp. that were placed on top of each other, as well as a bud of Quercus sp. and a leaf of Betula sp. The larva spun together a portion of two of the Quercus sp. leaves. On 13 July Sheldon provided it with a dead leaf of Fagus sylvatica for pupation and a few days later found that it had spun a cocoon within a small portion of that leaf, which it had folded over. The moth emerged on 15 May the following year. Sheldon does not explain why he provided the larva with a dead leaf of Fagus sylvatica when it had been feeding on leaves of Quercus sp., but the fact that he mentions that this leaf was dead and makes no such comment about the other leaves presumably implies that those were fresh. He describes three instars which we set out in full as they are the most comprehensive that have been published. First instar . ‘2 mm. long, whitish-green in colour, the head was glabrous, and dark fuscous; the prothorax was whitish-green with a dark fuscous line at the rear, and in the centre of the dorsum; the larva was very spiny.’ Second instar . ‘2.50 mm. long; the head and prothorax were amber-coloured; both were glabrous. The segments behind the prothorax were dark greyish-green in the dorsal, and light grey in the spiracular areas; the larva was very transparent.’ Third instar . ‘5 mm. long, slender, the head was amber-coloured with dark brown shading around jaws, glabrous and transparent; at the rear it had two distinct lobes. The rear half of the prothorax was the same colour as the head; the front half was much paler. Next the mesothorax are two half-crescents, dark brown in colour, divided in the centre by a thin line of light colour. The segments at the rear of the prothorax are light honey-coloured and very transparent. There is an unusual arrangement of colour in the dorsal regions; this includes the whole of those from and including the rear portion of the prothorax, and all the segments at the rear of it with the exception of the last four. This colour is dark brown, and it is continued to its anal extremity of the same width. The tubercles [pinacula] are slightly lighter than the surroundings, but are not prominent. The spiracles are slightly darker than the adjoining area; they are not conspicuous. The claspers and prolegs are dark fuscous. The alimentary canal shows as a distinct dark line through the brown dorsal area, but is hardly visible at the rear of it. The anal plate is not noticeable.’ Ent Rec 131(3).qxp_Layout 1 25/05/2019 10:19 Page 114

114 Entomologist’s Rec. J. Var. 131 (2019) The description of the third instar was made on 7 July. Sheldon states that on 13 July ‘the larva was 9 mm. long, apparently still in the third instar; it was now very similar to when last described, with the exception that the anterior segments were now almost as dark as those in front of them.’ He then records, as mentioned earlier in this account of his observations, that on the same day he gave it a dead leaf of Fagus sylvatica and a few days later found that it had spun a cocoon. Accordingly, either that larva underwent only three instars or Sheldon did not observe at least one of them. In the same volume as Sheldon’s account of the larva, Thurnall (1920) provides a note of his discovery of the larva. This is much shorter and less detailed than Sheldon’s and reads as follows: ‘early in July [?1916] I was collecting near Brentwood, and from a shrubby oak [ Quercus sp.] I beat out two little dark-coloured, extremely active larvae that were quite strangers to me. Upon examining them carefully at home I came to the conclusion that they must be larvae of L. permixtana , the moth being common there. Both were full fed and spun up the next day, turning a lobe of an oak leaf over something like an Ornix but not so flat. On May 31st, 1917, a male appeared, and upon examining the other one I found it had pupated but was dead. I think oak must be its favourite pabulum in this part of the country, but I have certainly beaten the imago from birch [ Betula sp.] as well.’ Swatschek (1958: 195) describes the larva, in translation, as ‘brownish yellow, with red longitudinal stripes dorsally or dull brown-red. The body is strongly granulated dorsally by small setae. The pinacula are pale, sometimes brown on the thorax, the head, prothoracic and anal plates light brown, the prothoracic plate sometimes edged darker’. He states that the description is based on preserved larvae deposited in the Zoologische Staatssammlung, München, Germany, and found by Disqué on 1 October 1917 at Speyer (in the Rhine valley, Germany), ‘im umgeschlagenen Blatt an’ (‘in the folded leaf of’) Betula (species not stated). He gives the larval period as ‘VII, V und IX’ with the following foodplants: in the tips of the stem of Solidago virgaurea and Anchusa officinalis as well as Betula and Fagus (species not stated for either). The first part of the account, with the mention of red longitudinal dorsal stripes, does not accord with our observations or those of Wood (1888) or Sheldon (1920), or the two images of one larva on Lepiforum (2019). This part of the description has, however, apparently been followed by Hannemann (1961: 199) and in part by Bradley, Tremewan & Smith (1979: 68) and Hancock & Bland (2014: 21– 22). Razowski (2003: 23) appears to condense Swatschek’s alternative descriptions into one, giving the larva as brownish yellow with greenish or reddish admixtures, with a reddish brown dorsal line (not lines or stripes as given by Swatschek). The alternative colour of the body, dull brown-red, given Ent Rec 131(3).qxp_Layout 1 25/05/2019 10:19 Page 115

Entomologist’s Rec. J. Var. 131 (2019) 115 by Swatschek ( loc. cit .) could apply to Lobesia reliquana . The date, 1 October 1917, would seem to be late for the larval period. Further, there is no indication that Disqué retained one or more larvae and reared moths. Therefore it is unclear to us on what basis he determined the larvae that he preserved as L. reliquana . Hannemann (1961: 199) describes the larva as ‘bräunlichgelb mit rötlichen Dorsalstreifen, Kopf und Nackenschild hellbraun’ (b rownish yellow with reddish dorsal streaks, head and prothoracic plate light brown). This appears to be an abbreviated version of the account by Swatschek (1958: 195). Bradley, Tremewan & Smith (1979: 68) describe the larva as follows: ‘Head and prothoracic plate shining light brown; thoracic legs black; abdomen with segments deeply incised, varying from brownish yellow, sometimes tinged with green with a reddish brown dorsal stripe, to dull purplish red; pinacula paler than integument, small, inconspicuous; peritreme of spiracles blackish; anal plate inconspicuous (Wood, 1888: 66).’ What is curious about this is that although their description is based on Wood’s 1888 account, there are various significant differences. Wood does not describe the abdomen as ‘varying from brownish yellow, sometimes tinged with green with a reddish brown dorsal stripe, to dull purplish red’, but simply as ‘dark puce’. ‘Puce’ is no longer a word used in the British literature when describing larval colour, but means dark red or purple-brown. It is possible that Bradley, Tremewan & Smith were following Swatschek (1958: 195) at least in part. Moreover, although Wood states that the pinacula (spots) are small, he notes that they are conspicuous and white whereas Bradley, Tremewan & Smith describe them simply as paler than the integument and inconspicuous. Our observations, set out more fully later, in general agree with Wood’s. Finally, although Bradley, Tremewan & Smith give the peritreme of the spiracles as blackish, Wood makes no mention of them. Our observations agree with Bradley, Tremewan & Smith on this point. Razowski (2003: 23) provides the following larval description stated to be after Bradley, Tremewan & Smith, but which differs in a significant respect as regards the body colour. ‘Larva varies in colour from brownish yellow with greenish or reddish admixtures, with reddish brown dorsal line; pinacula small, paler than body; stigmata edged black; head and prothoracic shield glossy light brown; thoracic legs black; anal shield indistinct.’ The result is that according to this wording the body always has a reddish brown dorsal line whereas Bradley, Tremewan & Smith describe the abdomen as varying from brownish yellow to dull purplish red, but it is only the former colour that is stated to be sometimes tinged with green and with a reddish brown dorsal stripe. Ent Rec 131(3).qxp_Layout 1 25/05/2019 10:19 Page 116

116 Entomologist’s Rec. J. Var. 131 (2019) Hancock & Bland (2014: 21–22), in The Moths and Butterflies of Great Britain and Ireland volume 5 (2), give the same larval description as Bradley, Tremewan & Smith (although parts are in a different order), also attributing this to Wood (1888). Lepiforum (2019) has two images of the same larva. This has the head honey- brown and shiny, the prothoracic plate darker honey-brown, edged blackish brown posteriorly and laterally, the body almost translucent greyish brown, slightly shiny with very pale yellowish white pinacula anteriorly becoming indistinct posteriorly bearing short white setae, and the anal plate concolorous with the body.

Our observations on the larva On 12 July 2009 we found three similar larvae, one amongst a withered brown leaf of Carpinus betulus and two amongst withered brown leaves of Quercus sp., at Budshead Wood, Devon. This is a small wood mainly comprising mature Carpinus betulus and Quercus sp. with some Betula sp. We suspected that the larvae might be those of Lobesia reliquana and confirmation was provided when a moth emerged on 2 May the following year. This resulted from one of the larvae that was amongst the Quercus sp. leaves. Unfortunately that was the only moth that we reared. On 9 August 2010 we found several similar larvae at the same locality again amongst withered brown leaves of Carpinus betulus and Quercus sp. Those amongst the latter were often between two spun leaves. We failed to rear moths. On 20 August 2013 we found one similar larva at Hembury Woods, Devon amongst withered brown leaves of Quercus sp. but failed to rear the moth. On 3 September 2013 we found three similar larvae again at Budshead Wood, all amongst withered brown leaves of Quercus sp. From these one Lobesia reliquana resulted on 31 March 2014. All the leaves were attached to twigs caught in the lower branches of trees or bushes. The larvae were within silken spinnings and, as noted by Wood (1888) and Thurnall (1920), were very active when disturbed. We did not specifically observe when the larvae ceased feeding, but it was in the early autumn when cocoons were spun often amongst the withered leaves and sometimes amongst the tissue in the containers in which the larvae had been kept. We did not note whether the larvae folded down part of the edges of the leaves within which to pupate, as by that stage the leaves were too withered to investigate.

Description Larva . Final instar . (Plate 1). Head honey-brown, shiny; prothoracic plate darker honey-brown, suffused blackish brown posteriorly and laterally or edged Ent Rec 131(3).qxp_Layout 1 25/05/2019 10:19 Page 117

Entomologist’s Rec. J. Var. 131 (2019) 117

Plate 1 . Larva of Lobesia reliquana (Hübner, [1825]). England: Devon, Budshead Wood, larva found within withered brown leaf of Quercus sp. 3.ix.2013, photographed 4.ix.2013 at 12.41, moth emerged 30.iii.2014. Photo : R. J. Heckford

Plate 2 . Larva of Lobesia reliquana (Hübner, [1825]). England: Devon, Budshead Wood, larva found within withered brown leaf of Quercus sp. 3.ix.2013, same larva as in Plate 1 photographed 4.ix.2013 at 12.43, moth emerged 30.iii.2014. Photo : R. J. Heckford Ent Rec 131(3).qxp_Layout 1 25/05/2019 10:19 Page 118

118 Entomologist’s Rec. J. Var. 131 (2019) black posteriorly and laterally; body reddish brown, body contents sometimes showing dark grey; pinacula white, comparatively large and conspicuous on thoracic segments 2–3, greyish white, smaller and sometimes less conspicuous on abdominal segments 1–8, slightly larger on abdominal segment 9; setae white; peritremes of spiracles blackish; anal plate large, paler than body, grey with black marks; thoracic legs black; ventral and anal prolegs whitish, crochets unobserved. Depending on the substrate the colour of the body of the larva can appear to change, but this is an artefact. Plate 2 shows the same larva as in Plate 1 photographed with the same equipment used for Plate 1 only two minutes later. Pupa. Not described, in a cocoon. Exuviae pale yellowish brown, not extruded on emergence of adult.

Hymenopterous parasitoids reared from larvae of Lobesia reliquana As well as rearing two moths, we also reared two Hymenopterous parasitoids. Choeras ?tedellae (Nixon, 1961) (Braconidae, Microgastrinae), 1 & reared on 19 August 2010 from a larva collected at Budshead Wood, Devon amongst withered leaves of a Quercus sp. on 9 August 2010. Although the specimen appears to be structurally identical with Choeras tedellae it is much darker than normal specimens and the host is surprising, because it is otherwise only known from Epinotia tedella (Clerck, 1759) (Lepidoptera: Tortricidae, Olethreutinae) whose larva feeds on Picea abies (L.) H. Karst. and Picea sitchensis (Bong.) (Carrière). The identity of this specimen is discussed by Shaw (2012: 176). Phytodietus sp. (Ichneumonidae, Tryphoninae), 1 & reared on 1 October 2013 from a larva collected at Budshead Wood amongst withered leaves of Quercus sp. on 3 September 2013.

Summary Various questions arise from the records cited earlier in this paper as to what are the true larval foodplants of Lobesia reliquana . It is clear that the larva of Lobesia reliquana is polyphagous, with 18 foodplants being cited in this paper. However, for the reasons given we consider that the following eight of these, in the order in which they were published, are either incorrect or require confirmation: Juniperus communis , Anchusa officinalis, Solidago virgaurea , Fagus sp., Ulmus sp., Corylus avellana , Salix sp. and Alnus glutinosa . The following 10 are definite foodplants, again in the order in which they were published except the last two which appear on Lepiforum (2019) but may have been placed there much earlier as the larva, pupae and pupation sites whose images appear there were observed between 2008 and 2009: Prunus spinosa , Betula pubescens , Alnus incana , Quercus sp., Ent Rec 131(3).qxp_Layout 1 25/05/2019 10:19 Page 119

Entomologist’s Rec. J. Var. 131 (2019) 119 Betula pendula , Prunus avium , Carpinus betulus , Rubus ulmifolius , Quercus robur and Cornus sanguinea . As regards larval descriptions, those of Wood (1888) and Sheldon (1920) are undoubtedly reliable, being based on larvae from which moths resulted. That of Kennel (1910: 272) agrees with those and in general with our observations, and the two images of the larva on Lepiforum (2019), even though the five foodplants that he cites are either incorrect or require confirmation. Other publications that we have cited have parts that agree with undoubted larval descriptions and parts that are at variance. We suggest that any larva found in the wild thought to be Lobesia reliquana should be reared unless it completely agrees with the descriptions that we consider to be reliable or with the images on Lepiforum and in Plates 1 and 2. Also, any larva potentially of this species that is found on any of the eight foodplants that we consider doubtful or incorrect should also be reared, even if it agrees with these descriptions and images. Finally, perhaps any larva potentially of this species should be reared, irrespective of whether it is on a known foodplant in case it might prove to be Lobesia virulenta mieana which would be new to Britain!

Acknowledgements We are extremely grateful to Dr K. Sattler (Natural History Museum (BMNH), London) for his assistance with the translation and understanding of certain German texts and Dr M. R. Shaw (National Museums of Scotland, Edinburgh) for identifying the parasitoids to genus level. We thank Dr J. R. Langmaid (Southsea) for allowing us to publish his observations on the circumstances of where he found overwintering larvae. We are also indebted to Mr J. Nunes (Valongo) for providing us with more information about the larval foodplant than given in Corley et al . (2018) as well as his excellent unpublished images of the larva and larval pupation site, and to Mr M. V. F. Corley (Faringdon) for putting us in contact with him.

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