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MONOPIS CROCICAPITELLA (CLEMENS, 1859) (LEP.: ): CASE-BEARING LARVAE IN ENGLAND FOUND FEEDING ON BAT DROPPINGS R. J. H ECKFORD ¹ AND S. D. B EAVAN ² ¹ Department of Life Sciences, Division of , Natural History Museum, Cromwell Road, London SW7 5BD (Correspondence address: 67 Newnham Road, Plympton, Plymouth, Devon PL7 4AW) ² The Hayes, Zeal Monachorum, Devon EX17 6DF

Abstract An account is given, including a larval description, of crocicapitella (Clemens, 1859) being reared from case-bearing larvae found feeding on bat droppings in Devon, England. Although there have been three notes in the British literature recording the as a case-bearer, these appear to have been overlooked in subsequent publications. There has been no prior report of the larva feeding on bat droppings in the British Isles, nor any larval description based on British observations. We consider records made both within and outside the British Isles. Key words : , Tineidae, , larval case, larva, description, bat droppings.

Introduction On 11 September 2017 Dr M. L. Luff made one of his frequent visits to Dartington Hall Estate, Devon, England, for the purpose of studying Coleoptera. During the course of this, and in the company of Mr C. N. Wills, Site Bat Warden for the Devon Bat Group, he collected a quantity of bat droppings that had been produced by Lesser Horseshoe Bat Rhinolophus hipposideros (Bechstein, 1800) (Chiroptera), in the hope that they might harbour beetles, either adults or larvae. Unfortunately they did not but he noticed a number of larval cases which he thought might be of interest and the following month passed these to us together with the bat droppings. Later that month began to emerge and proved to be Monopis crocicapitella (Clemens, 1859). This was quite a surprise for two reasons. One was because we were unaware that the larva made a case and the other was because we did not know that it fed on bat droppings. Pelham-Clinton (1985: 189), in a comprehensive review of British Tineidae in The Moths and Butterflies of Great Britain and Ireland 2, states that the larva is apparently undescribed, although it had been reported to feed on a variety of materials which are listed but bat droppings are not mentioned. Moreover, later in that volume Pelham-Clinton (1985: 195) comments that the larvae of most in the Linnaeus, 1758, feed from a portable case, whereas there is no suggestion in his account of the genus Monopis Hübner, [1825], that the larva of any species in that genus does so. Ent Rec 130(5).qxp_Layout 1 14/10/2018 11:02 Page 234

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In fact Bignell (1878) had recorded that the larva of Monopis crocicapitella was a case-bearer, although his note is entitled ‘Food of Tinea ferruginella ’; Tinea ferruginella is now ([Denis & Schiffermüller], 1775). In 1878 Monopis crocicapitella had not been recognised as a British species, no doubt because the adult looks similar to M. obviella . M. crocicapitella was tentatively added to the British list by Richardson (1893) who described it as a new species under the name Blabophanes heringi . Walsingham (1907: 728) synonymized this with Monopis crocicapitella and subsequently Bankes (1912) stated that N. M. Richardson had ascertained that Bignell’s material was in fact Monopis crocicapitella . Bignell did not describe either the larva or the case and as far as we are aware the first description of both was provided by Căpuşe & Georgescu (1963) in a Romanian journal. There are, however, other publications, both in Britain and elsewhere, which refer to the larva being a case-bearer. Some also either state that the larva feeds on bat droppings or associate the species with bats in caves. We are not aware, however, of any larval description or records of the larva feeding on bat droppings based on British observations. Therefore later in this paper we describe, and illustrate, the larva and its case but first we review certain published records of the biology made in the British Isles and abroad.

Observations on the larva made in the British Isles Bignell (1878) was apparently not only the first in the British Isles to rear the species, but possibly the first person anywhere to do this. Although he gives no larval description, importantly he refers to the fact that the larva is a case-bearer. In August 1876 he gathered a quantity of Wormwood Artemisia absinthium L., locality not stated, in order to obtain certain Macrolepidoptera larvae. He placed this in a linen bag which was hung up, presumably outside. On 21 February 1877 he searched the contents and about half a dozen ‘ Tinea ferruginella ’ flew out which he captured. As recorded in the Introduction to this paper, Bankes (1912) states that subsequently these were identified as Monopis crocicapitella. Bignell comments that ‘I then collected all the cases I could find; some were in pupa, others in the larva-state and feeding.’ Bignell sent some cases to C. G. Barrett and records that his reply was that ‘it was a welcome and startling sight to find it cleverly clearing out the seeds from every flower-head of the dry wormwood.’ Bignell concluded that the problem of what the larva eats had now been solved, as it was a seed-feeder, but wondered whether it was confined to species of Artemisia or whether it also fed on other ‘composite plants’ (namely species in the Compositae, now usually given as the Asteraceae), because the was known to occur where Artemisia absinthium did not. Meyrick ([1928]: 823) gives ‘Larva on seeds, woollen refuse, also probably on dried excrement, etc.’ It is likely that some of this information came from Ent Rec 130(5).qxp_Layout 1 14/10/2018 11:02 Page 235

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sources outside the British Isles. He makes no mention of the larva inhabiting a case unlike, for example, his account of Tinea pellionella Linnaeus, 1758, at page 830. Ford (1949: 180) states, ‘On seeds, woollen refuse, refuse of birds’ nests.’ Like Meyrick ( loc. cit .) he makes no mention of the larva inhabiting a case but does so in his account of Tinea pellionella at page 184. Then Woodroffe & Southgate (1952) record the larva as being a case-bearer. This was as a result of the discovery of a considerable infestation in a block of flats at Harrow, Middlesex. A hot-water pipe was lagged along much of its length with asbestos, but beneath two flats the lagging was of felt which was covered with case-bearing larvae and in one flat the larvae were attacking the underfelt of the hall carpet. Rather surprisingly, Woodroffe & Southgate do not describe either the larva or case. Obviously unaware of Bignell’s observations, they comment that ‘the literature appears to contain no reference to the fact that the larvae are case-bearers, resembling larvae of Tinea pellionella L.’ The following year Woodroffe (1953: 745) states that ‘The larvae are case- bearers, and were found several times in some numbers in pigeons’ nests.’ In the previous sentence, however, he refers to his note jointly published with Southgate and cited in the above paragraph, and so his comment that the larvae are case-bearers does not necessarily mean that those found in pigeons’ nests were also case-bearers. Unfortunately Robinson (1979), in a detailed account of what is ‘a revision the case-making (or ‘case-bearing’) clothes-moths of the Tinea pellionella group’, was obviously unaware of the notes published by Bignell, Woodroffe & Southgate and Woodroffe because at page 58 he comments that: ‘Only a few other species of Tineidae with case-making larvae are occasionally found associated with woollen or feather products – these are free-living Monopis species and their larval ‘cases’ are not true cases but immovable sections of tunnel, the walls of which consist predominantly of frass and food particles.’ In the following year, however, Robinson (1980: 107) states that ‘Rather surprisingly, the only description of the larva of this common and widespread species seems to be that by Căpuşe & Georgescu (1963b: figs. 1–7). The larva makes a flattened, slightly ovate, fine-grained case.’ The reference to Căpuşe & Georgescu (1963b) is to their paper cited both in the Introduction to this account and later. Pelham-Clinton (1985: 189), a meticulous observer, states that the larva is apparently undescribed, and with no suggestion that the larva is a case-bearer. He records that the larva is ‘reported as feeding on a variety of materials including flour, oats, other seeds and woollen refuse’ and comments that moths are ‘sometimes common in poultry-houses and have been bred from pigeons’ Ent Rec 130(5).qxp_Layout 1 14/10/2018 11:03 Page 236

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nests.’ Although he was correct in commenting that the larva was apparently undescribed as far as the British literature was concerned, nevertheless, as already mentioned, Căpuşe & Georgescu (1963) had already described the larva, and larval case and pupa, which we consider in the next section. Later Pelham-Clinton (1988: 46) gives the larva as ‘In birds’ nests, on stored products of vegetable origin including flour, corn, felt and , or on a variety of refuse of plant or origin.’ Again there is no mention that the larva is a case-bearer, in contrast to the comment on the following page that, with the exception of Tinea pallescentella Stainton, 1851, so far as is known all the species in the genus Tinea construct portable silken cases. The purpose of the work was to enable a field-worker to recognise a species from its field characteristics and so there are no descriptions of larvae, or adults. Sterling & Parsons (2012: 77) give the pabula as ‘Refuse of plant or animal origin, including birds’ nests’, with no mention of the larva being a case-bearer. This is in contrast to their account of Tinea pellionella on the previous page where they illustrate cases and state that the larva lives in a portable case. Manley (2015: 40) gives ‘Birds’ nests, plant and animal refuse’ as the pabula. He makes no mention of a case, but this might not be significant because he does not state that Tinea pellionella , which is on the same page, makes a case, although one with exuviae is illustrated on the facing page. The latest account of the larva in the British literature is provided by Boyes (2018: 59) and, like Pelham-Clinton (1988: 46), he makes no mention of the larva being a case-bearer or feeding on bat droppings. In summary, the fact that the larva of Monopis crocicapitella makes a case has been mentioned three times in the British literature: by Bignell (1878), Woodroffe & Southgate (1952) and Woodroffe (1952: 745). These publications, however, appear to have been overlooked until now. Finally, there appears to be no mention that the larva will feed on bats’ droppings or that, according to certain observations including our own, damp conditions may be a larval requirement, which we discuss later.

Observations on the larva made outside the British Isles There are several accounts outside the British Isles of the larva of Monopis crocicapitella being a case-bearer, and a few of the species being associated with bat caves and of the larva feeding on bat droppings. When Clemens (1859: 258) described Monopis crocicapitella , as Tinea crocicapitella , he made no mention of the larva. Forty years later Hering (1889: 295–299) described Blabophanes lombardica sp.n., subsequently synonymized by Walsingham (1907: 728) with Monopis crocicapitella . He records that the larva is case-bearing. Although he comments Ent Rec 130(5).qxp_Layout 1 14/10/2018 11:03 Page 237

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that the description of the larva must be postponed until later (‘ Die Beschreibung der Raupe muß auf später verschoben werden’) , he gives the following account of the case: ‘Der Sack, welchen ich in mehreren Ex. mit dem Falter erhielt ähnelt durchaus dem von T. pellionella L., columbariella Martini und ähnlichen. Er ist etwa 8 mm lang, flach, schmutzig-erdgrau, das Analende mit weicheren Wolltheilen gefasert, sonst mit einzelenen Sandkörnen und Kothklümpchen bedeckt. Er wird vermuthlich nach den verschiedenartigen Fraßsubstanzen auch verschieden in Farbe und Form ausfallen. Die mattgelbbraune Puppenhülse dringt zum Theil nur bis zur Hälfte der Flügelscheiden, zum Theil bis zum 4. Segment aus dem Sack hervor.’ (The case of which I received several examples with the moth resembles that of T. pellionella L., columbariella Martini and the like. It is about 8 mm long, flat, dirty-earth grey, the anal end with softer wool parts, otherwise covered with individual grains of sand and frass pellets. It will probably vary in colour and shape depending on the various feeding substances. The dull yellow- brown pupae penetrate out of the case partly up to the middle of the wing, partly up to the fourth segment.) The first reference that we can trace to the species being associated with bats is given by Forbes (1923: 132). He states that ‘The larva has been bred from seeds of absinth and refuse, and the moth has been taken in a cave, associated with bats.’ We presume that the reference to absinth was taken from Bignell (1878). Hindwood (1951: 129–130) records larvae being found in a nest of Fairy Martin Hylochelidon ariel (Gould, 1843), now Petrochelidon ariel , and apparently feeding on feathers. He also refers to pellets produced by both Little Eagle Hieraaetus morphnoides (Gould, 1841) and Barn Owl Tyto alba (Scopoli, 1769) being infested with larvae from which Monopis crocicapitella was reared. He makes no mention of larval cases. As far as we can trace, and in agreement with Robinson (1980: 107), the first description of the larva was provided by Căpuşe & Georgescu (1963) who also describe the larval case and pupa, all illustrated in a monochrome plate including an image of the larval case with extruded exuviae, together with figures of the chaetotaxy of the thoracic and abdominal segments. The larvae were found by M. Dumitrescu and T. Orghidan during speleological research in the Gura-Dobrogei cave in Romania amongst bat guano. Căpuşe & Georgescu provide the following larval description: ‘Larva (fig. 7). Lungimea corpului 10 –12 mm. Lăţimea 1,8 –2 mm. Culoarea capului, a plăcilor protoracelui, a plăcii anale, picioarele şi peritrema stigmelor brună. Restul cuticulei de culoare albă-gălbuie-murdar şi acoperită cu microtrichi deşi în formă de spini aciculari. Ent Rec 130(5).qxp_Layout 1 14/10/2018 11:03 Page 238

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Capul , lipsit de oceli, prezintă lateral o bandă lată brună-negricioasă care se întinde pînă în apropierea setei O 2. Stigmele rotunjite. Mandibulele bine dezvoltate prezintă 4–5 dinţi puternici, iar pe marinea externă, 2 sete. Labrumul are incisura puţin adîncă, iar cea mai lungă setă este L 2. ’ Dr M. Stănescu, Collection Manager, Lepidoptera, The “Grigore Antipa” National Museum of Natural History, Bucharest, Romania, tells us that the text is cumbersome even for a Romanian reader. His following translation has been slightly amended by us. ‘Larva (fig. 7). Body length 10–12 mm. Width 1.8 –2 mm. Head, prothoracic plate, anal plate, legs and peritremes of spiracles brown. The rest of the cuticle dirty yellowish white covered with microtrichia, although in the form of acicular thorns. Head , without ocelli, shows on each of its sides a broad lateral blackish brown stripe which extends up to the O 2 seta. Spiracles rounded. Well-developed mandibles with 4–5 strong teeth, on their outer edge with two setae. Labrum has a shallow incision, the longest seta being L 2.’ The word ‘ picioarele’ in the first paragraph of the original description means ‘legs’, but this word does not differentiate between the thoracic legs and the prolegs, and Căpușe & Georgescu do not make clear to which they are referring, or whether they are referring to both. They then give an extremely detailed account of the chaetotaxy, which Dr Stănescu also kindly translated for us, but which we do not set out as it needs to be read in conjunction with the figures of the thoracic and abdominal segments included in their paper . As regards the larval case this is described as being compact, transportable and has the appearance of a glasses case open at both ends, but no mention is made of its colour or from what it is constructed. The two images of the case, however, one with exuviae extruded to about the fifth abdominal segment, clearly show that it has very small particles attached and is dark. A detailed description is provided of the pupal structure including the observation that t he 3rd to 8th abdominal segments are mobile and provided dorsally with a line of strong spines directed backwards and that the cremaster bears 2 lateral grooves (‘ Segmentele abdominale de la 3-lea pînă la al 8-lea sînt mobile şi prevăzute dorsal cu un şir de spini puternici îndreptaţi posterior. Cremasterul poartă 2 excrescenţe laterale.’) No indication is given of the colour of the pupa. A Hymenopterous parasitoid was reared: Hemiteles flavigaster Schmiedekecht, 1897 (Ichneumonidae, Cryptinae), now a junior of Charitopes gastricus (Holmgren, 1868). A few years later Căpuşe (1968: 370–374, figs 186, 192 and 193) gives a fuller account of the species including descriptions of the adult and the genitalia Ent Rec 130(5).qxp_Layout 1 14/10/2018 11:03 Page 239

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of both sexes. There is a monochrome plate with the same images of the larva, larval case, larval case with extruded exuviae and pupa as in the 1963 joint publication with Georgescu, but with the addition of an image of a set adult. The larval description is in almost exactly the same terms as in the 1963 account, including the same figures of the chaetotaxy. The larval case is stated to be made from sand, dust and guano (‘construite din nisip, praf şi guano’). The detailed pupal description is in almost exactly the same terms as that in 1963, and again no indication is given of the colour. As regards the biology, it is clearly stated that, amongst other things, the larva feeds on the excrement of raptors and bats ( şi se hrănesc ... excremente de păsări răpitoare şi lilieci.’) Zimmerman (1978: 320) records that ‘The larvae have been found in debris, in a dead rat, and in “ stuffs”, and Don R. Davis has recently found the larvae associated with pigeon guano on Hawaii.’ There is no suggestion that the larva makes a case, in contrast to his earlier account (Zimmerman, 1978: 278) of the larva of Tinea pellionella that it ‘constructs and always bears a case’. Carter (1984) did not treat the larva as a case-bearer, despite the fact that he provides a larval description based on Căpuşe & Georgescu (1963). In the first couplet of a key to larvae in the Tineidae of economic importance in (Carter, 1984: 20) he divides the larvae into case-bearing and non case-bearing, and the latter couplet eventually leads to ‘ Monopis spp.’ The account of Monopis crocicapitella appears at pages 51–52. There the following larval and pupal descriptions are given: ‘Larva . Head brown with a broad blackish brown stripe along each side behind ocelli; body dirty yellowish white; peritremes of spiracles, thoracic legs, prothoracic and anal plates brown (Căpuşe & Georgescu, 1963). Pupa . Yellowish brown; abdominal segments 3–8 each with one transverse dorsal band of spines; cremaster with two ventrolateral, conical processes and a dorsal process with two bifid prongs.’ Although Carter specifically describes the thoracic legs as brown, as stated earlier in this section the word used by Căpuşe & Georgescu, ‘picioarele ’, does not differentiate between thoracic legs and prolegs. Later in his account Carter states that the larva occurs from August–April in birds’ nests or bat roosts, and the pupa from May–July ‘in a flattened cocoon of sand and fine debris.’ It is clear that Carter does not consider this cocoon to be a case because at the end of his description of the larva of Tinea pellionella he states, ‘In a fusiform, dorsoventrally flattened case’. We do not know why Carter was apparently unaware that the larva could be a case-bearer in view of his larval description being based on Căpuşe & Georgescu. Robinson & Nielsen (1993: 167–168) state that the larvae of the genus Monopis are keratophagous and chitinophagous and feed on a wide variety of substances: feathers, fur, wool, hides, guano in birds’ nests, and on animal Ent Rec 130(5).qxp_Layout 1 14/10/2018 11:03 Page 241

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remains in general. As regards the larva of M. crocicapitella they comment that in Australia it has been found feeding on guano in a number of different caves inhabited by bats. They record that they had seen a dense domestic infestation in a slightly damp woollen carpet, containing some fungal hyphae, beneath a wicker basket and provide the following account: ‘Larvae were in flattened, slightly ovate portable cases, pupae in similar but rougher cases firmly attached horizontally to the carpet or basket above (unlike Tinea spp. which attach the mouth of the case to the substrate on pupation). Pupae were almost completely extruded from the case, the exuviae remaining at an angle of about 45° to the case following emergence.’ Gerard (1995) also reports severe larval damage to a carpet in New Zealand. As a result of rearing adults and providing them with fresh strands of yeast- treated wool, ova were freely laid. She was therefore able to describe both the ovum and larva. She states that ‘The eggs were translucent white, oval in shape, slightly broader at one end than the other, with a dimpled surface.’ They were laid singly, loosely adhered to the substrate and had sufficient resilience to be dislodged with a paint brush without damage. The account of the larva is the most detailed that we have traced that is in the English language and so we set it out: ‘All larval stages had a tan head capsule and glossy creamy-white body. The prothoracic shield had two triangular pale amber plates on the dorsal surface, which allowed the larvae to be differentiated easily from Tinea dubiella [Tinea dubiella Stainton, 1859] larvae which have darker, more rectangular grey-brown plates. Neonate first instar larvae did not form webbing, but towards the end of that stadium, light webbing tunnels were found amongst the wool fibres. From the second instar onwards, larvae were usually found in a dorso-ventrally flattened portable case open at both ends. The case was covered with frass and wool laid down in an oval alignment. Following moulting, the shed head capsule was usually lightly attached to the outer case surface. With the addition of further case layers, the shed head capsules were found to be completely incorporated in the case matrix. The mature larvae reached about 10 mm in length, with cases 9–12 mm in length and 2.5–3.5 mm in breadth. In contrast to dubiella , disturbed M. crocicapitella larvae were found much more likely to abandon their cases than to retreat inside. In culture, M. crocicapitella was found to pupate within the larval case, sealing off both ends. The cases in which pupae occurred were frequently found firmly attached to the walls and lid of the rearing container by silk spun from both ends of the case.’ Figure 2 in Gerard’s paper is a monochrome photograph of a larval case with a larva extruded from it to about the first abdominal segment. Possibly because the figure is in monochrome, the head appears to be black and not ‘tan’. Ent Rec 130(5).qxp_Layout 1 14/10/2018 11:03 Page 242

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Interestingly, Gerard records that humidity is a relevant factor in the larval development. Optimum development occurred at 93% relative humidity with no survival at 25% and 51%. At 97% several larvae did not construct cases and the cases that were made were smaller than those made at 93%. According to the observations made, the larva has five instars. Tunca et al . (2014) deal with the rearing, in laboratory conditions, of a parasitoid from larvae of M. crocicapitella . Their paper does not describe the larva or mention that it is a case-bearer, but one of the colour photographs accompanying the paper shows three larval cases, two with larvae partially extruded. Eberhard et al . (2014: 41–42), in a paper on the invertebrate cave fauna of Jenolan in Australia, state that ‘ The guanophilic tineid moths Monopis crocicapitella (Clemens, 1859) and Hofmannophila pseudosprettella [sic ] (Stainton, 1849) have been reported from within the caves associated with bat guano.’ They refer to an Australian paper published in 1967 that commented that ‘ Monopis sp. moths have been ‘found in almost all bat-inhabited caves of eastern Australia, where the larvae develop on heaps of guano’.

Our observations in 2017 and 2018 As mentioned in the Introduction, on 11 September 2017 Dr M. L. Luff, in the company of Mr C. N. Wills, collected a quantity of bat droppings at Dartington Hall Estate, Devon that had larval cases which he kindly passed to us together with a quantity of the bat droppings. We were quite puzzled as to what they might be because they looked like cases made by a member of the Tinea genus but we were not aware of any member of that genus feeding on such pabulum in Britain. The mystery was resolved when the first Monoposis crocicapitella emerged the following month, and moths continued emerging until late February 2018. When we received the bat droppings they were fairly damp and the cases, all of which had larvae but no pupae, were either on the surface of the droppings (Plate 1) or slightly below the surface. All the cases were elongate, more or less rectangular, flattened dorso-ventrally and between about 8–10 mm long and about 4 mm wide. As both Căpuşe & Georgescu (1963: 833) and Gerard (1995: 57) record, the larval case is open at both ends. The larva can be quite active within the case because we observed larvae to partially emerge from one of the ends, retreat, turn around within their cases and emerge at the other ends, all within less than a minute. When the larvae are turning round the cases visibly bulge slightly. In locomotion the maximum extent that the larva emerges from its case is about up to the anterior part of the third abdominal segment (Plates 2, 3). Ent Rec 130(5).qxp_Layout 1 14/10/2018 11:03 Page 243

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On 25 January 2018 Dr Luff paid another visit to Dartington Hall Estate, again in the company of Mr Wills, and collected a further, large, quantity of droppings produced by Lesser Horseshoe Bat. These too had larval cases of M. crocicapitella which varied in size from about 3 mm long and 1 mm wide to about 10 mm long about 4 mm wide. Most of the larvae had not pupated. It was difficult to know whether some or all of these had been larvae at the time of the first visit on 11 September 2017 and had taken longer to feed or whether some, or all, had resulted from moths that had emerged sometime in the autumn/early winter and had mated. It certainly seems unlikely that the larvae in the very small cases, about 3 mm long, would have been feeding on the date of the first visit. Therefore either they were the result of ova laid before that date and had not hatched until fairly recently or were progeny from moths that had emerged since that date. We placed the larvae from both visits in several closed containers, all having the base lined with tissue. In some containers we placed only bat droppings, in others only birds’ feathers. The larvae fed on both pabula. Robinson & Nielsen (1993: 168) state that the larvae they observed were in slightly ovate cases and the pupae were ‘in similar but rougher cases’, thus suggesting that the larvae construct new cases within which to pupate. Our observations differ in two respects. The first is that, as mentioned earlier in this section, the cases we had were elongate, more or less rectangular and flattened dorso-ventrally. This accords with the photographs of the cases provided by Căpuşe & Georgescu (1963: 832), Gerard (1995: 56) and Tunca et al . (2014: 2074, fig. 1A). The second is that the larvae did not make fresh cases within which to pupate, nor is there any suggestion in any of the other literature that we have cited that the larvae do this. The larvae pupated either by fixing one end of the cases to the lids or sides of the containers so that they were suspended, or were more or less flat on the substrate. When suspended the larvae did not fix the other end of the cases to the lids or sides of the containers. When fixed more or less flat on the substrate, sometimes both ends were attached to this by silk and sometimes only one end was so attached. Robinson & Nielsen (1993: 168) appear to imply that the larvae pupate in cases that are attached at both ends to the substrate, but their observations were of cases fixed horizontally. Gerard (1995: 57) records larvae pupating in cases attached to the walls and lid of the rearing container by silk spun from both ends of the cases. Thus whether the cases are fixed by silk spun from both ends appears to be a variable factor. In general our observations on the colour of the larva are similar to the account given by Căpuşe & Georgescu (1963: 829) but differ in some respects. Rather than provide a complete description we set out where our observations differ from, or are additional to, that 1963 account. Ent Rec 130(5).qxp_Layout 1 14/10/2018 11:03 Page 244

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The head is reddish brown with black adfrontal sutures and does not have the broad lateral blackish brown stripe recorded by Căpuşe & Georgescu (1963). The colour of the prothoracic and anal plates, thoracic legs and peritremes of the spiracles is not the same as the head but is quite different. The prothoracic plate is slightly translucent and very pale yellowish brown with a narrow medial division of the body colour which is semi-translucent pale white, sometimes showing grey body contents. The anal plate is also very pale yellowish brown and almost translucent. The thoracic legs are semi-translucent pale white with small pale yellowish brown sclerotised areas on all segments except the coxa, the claw is darker yellowish brown. The peritremes of the spiracles are pale yellowish brown. Căpuşe & Georgescu (loc. cit. ) do not describe the colour of the pinacula, nor does Căpuşe (1968: 370–374). According to our observations the pinacula are semi-translucent and very pale yellowish brown or almost entirely translucent and colourless, and they are comparatively large. As mentioned in an earlier section, because Căpuşe & Georgescu use t he word ‘picioarele’, meaning ‘legs’, this does not differentiate between the thoracic legs and the prolegs, and so it is unclear whether their description that the legs are ‘brown’ refers to or includes the prolegs. Unfortunately we did not observe the latter. The only account that we can trace of the colour of the larval case is that by Hering (1889: 295–299) who describes it as dirty-earth grey, the anal end with softer wool parts, otherwise covered with individual grains of sand and frass pellets. He rightly surmises that it will probably vary in colour and shape depending on the various substances on which the larva feeds. There are several cases in the main collection of Lepidoptera in the Natural History Museum (BMNH), London. Some were collected by H. G. Amsel at Bremen, Germany. Each of these is pinned immediately below the stage-mounted adult. The data labels show that the moths emerged between November 1934 and January 1935 but do not record when the cases were collected nor on what the larvae were feeding. The cases are more or less rectangular and flattened dorso-ventrally. They have blackish fragments attached and do not show exuviae. We did not measure them but they appear to be about the same size as the largest cases from the bat droppings. In contrast there are cases collected in Canberra, Australia in July 1986. These are stage mounted separately from the resulting moths. They too are more or less rectangular but are more rounded dorso- verntrally. They are mainly grey and were probably constructed from man-made materials because some appear rather ‘woolly’ with fibres protruding although some also appear to have small black and white fragments attached. All have extruded exuviae, which extend far more than those that we observed from the cases found amongst the bat droppings, and one is almost at right angles to the case. So perhaps one or more of these cases came from the dense domestic Ent Rec 130(5).qxp_Layout 1 14/10/2018 11:03 Page 245

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infestation in a slightly damp woollen carpet in Australia recorded by Robinson & Nielsen (1993: 167–168), who state that the exuviae remain at angle of about 45˚ to the case. As regards our observations, the cases made by the larvae feeding on the bat droppings had dark blackish brown fragments attached to them which no doubt came from their pabula but probably also from their own frass, because we did not observe any frass when the larvae were in containers feeding on birds’ feathers. Occasionaly the cases also had shed head capsules affixed, as observed by Gerard (1995: 57). Sometimes the larvae in the containers with the birds’ feathers attached parts of these and if they were white these showed quite clearly. Our observations regarding the pupa agree with accounts that it is yellowish brown, with abdominal segments 3–8 each with one transverse dorsal band of spines directed backwards . The dorsal surface of many of the more primitive pupae often bears series of such spines to prevent the pupa from slipping back as it wriggles from its cocoon prior to ecdysis (Scoble, 1985: 132). Shortly before ecdysis the integument becomes translucent and the head, forewing markings and colours and body become clearly visible (Plate 4). There are differences in how far the exuviae extrudes from the case according both to published sources and our observations. Hering (1889: 295–299) records that the pupae penetrate out of the case partly up to the half of the wing and partly up to the fourth segment. Robinson & Nielsen (1993: 168) state that the ‘Pupae were almost completely extruded from the case’. As mentioned earlier in this section, some of the cases in the main Lepidoptera collection in the BMNH that were collected in July 1986 at Canberra, Australia, do have exuviae that are almost completely extruded, but only one is at an angle of 45° to the case. The exuviae from the cases found amongst the bat droppings are in almost the same plane as the cases and extrude no further than the third abdominal segment and about three-quarters the length of the wing case (Plate 5). The front of the head of the pupa is the sclerite to which the antennae are attached and this section of the pupa, together with the ventral section between the wing cases, is pushed forward at an angle of about 45° on emergence of the moth (Plate 6).

Discussion It is perhaps surprising that the publications by Bignell (1876), Woodroffe & Southgate (1952) and Woodroffe (1953) recording that the larva of Monopis crocicapitella is a case-bearer have remained overlooked in the British literature for so long, and that prior to 2017 there have apparently been no other observations of this in the British Isles. It may be, however, that the larvae are not easy to find. Hinton (1956) describes the larvae of 32 species of Tineidae Ent Rec 130(5).qxp_Layout 1 14/10/2018 11:03 Page 246

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from various areas of the world, including the British Isles. He states (Hinton, 1956: 252) that the larvae of most of the species known to be of economic importance had fortunately been made available to him, but he had not been able to obtain larvae of three species, one of which was M. crocicapitella . It is possibly less surprising that there have been no previous records of the larva feeding on bat droppings in the British Isles, because all bats are protected in the British Isles and licences are required if there is any possibility of disturbing them. Lesser Horseshoe Bats, like other members of the Chiroptera, are insectivores and their droppings consist largely of remains and therefore are suitable pabula. In general the droppings are brown to black, porous and crumble easily. They are often deposited in large heaps, which was the situation with the droppings found at Dartington Hall Estate. These were fairly damp when we received them and according to our observations the larvae required a damp environment in captivity. It was noticeable that they ceased feeding if it became too dry. This was easily seen if a larva was placed in a container with only a bird’s feather on the tissue lining the base of the sealed container. If the tissue was dry the larva would not feed but did so once the tissue was dampened. We did not attempt to measure the humidity, unlike Gerard (1995: 58–61) who records that optimum larval development occurred at 93% relative humidity with no survival at 25% and 51%. Robinson & Nielsen (1993: 167–168) had already observed in Australia a dense domestic infestation in a slightly damp woollen carpet, containing some fungal hyphae. Thus it seems that the larva requires a damp habitat and that the degree of humidity may be critical. Although examination of the literature shows that the larva of Monopis crocicapitella is known to be a case-bearer, it appears that it is a facultative case- maker. Hindwood (1951: 129–130) cites pellets produced by Little Eagle and Barn Owl as ‘being infested with larvae’. It is difficult to interpret this as meaning anything other than the larvae were within the pellets, and therefore not case-bearing. There are also records of larvae occurring in birds’ nests without any suggestion that they inhabit cases. Robinson & Nielsen (1989: 33) observe that some members of the Tineidae appear to be facultative case-makers citing observations on Monopis longella (Walker, 1863), a species known from Iran, Pakistan, India, China, Thailand, Vietnam, Korea, Japan and Russia (Far East) (Huang et al , 2011). Larvae of this species infesting an artificial birds’ nest fed either on the surface of the nest or tunnelled inside; those tunnelling were free-living but those on the outside made cases. Robinson & Nielsen ( loc. cit .) state that Monopis icterogastra (Zeller, 1852), an Australian species, together with an unnamed relative species, are also facultative case-makers. As regards Monopis crocicapitella , it appears that sometimes the larvae are case-bearing and sometimes not. The question is, what causes some larvae to become case- bearers not only in this species but also in at least three other Monopis species? Ent Rec 130(5).qxp_Layout 1 14/10/2018 11:03 Page 247

Entomologist’s Rec. J. Var. 130 (2018) 247 Acknowledgements We are very grateful to Dr M. L. Luff (Totnes) for providing us with the material which led to our observations and also to Mr C. N. Wills, Site Bat Warden for the Devon Bat Group, for facilitating the collection of the bat droppings and identifying the bat species. We also thank Mr J. Channon, Estate Manager, Dartington Hall Estate, for granting permission to Dr Luff to sample on the Estate and Dr K. Sattler (BMNH) for providing certain German translations which were considerably helpful. By coincidence a month or so before we received the bat droppings, Mr D. J. Mann, Head of Life Collections, Oxford University Museum of Natural History, contacted RJH about Monopis crocicapitella in another context and asked about images of larvae. Although at the time RJH was unable to supply any Darren Mann was able to provide two of the papers cited in this article, for which we are very grateful. Finally, our deepest thanks go to Dr B. Zlatkov, Institute of Biodiversity and Ecosystem Research, Bulgarian Academy of Sciences, Sofia, Bulgaria, for putting us in touch with Dr M. Stănescu, Collection Manager, Lepidoptera, The “Grigore Antipa” National Museum of Natural History, Bucharest, Romania, to whom we owe an enormous debt of gratitude for the considerable amount of Romanian translation that he provided for us. Even though we did not use all of it in this paper it was extremely helpful.

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