EUROPEAN JOURNAL OF ENTOMOLOGYISSN (online): 1802-8829
http://www.eje.cz

Eur. J. Entomol. 116: 442–449, 2019
doi: 10.14411/eje.2019.045
ORIGINAL ARTICLE

Differences in the niches of keratin/chitin feeding moths(Lepidoptera: Tineidae) in bird nests in central JapanHIROAKI SATO 1, YOSHITSUGU NASU 2, SHIRO MURAHAMA3, HIROYUKI MATSUMURO 3 and KEISUKE UEDA4
Department of Biological Sciences, Faculty of Science, Nara Women’s University, Kitauoya-Nishimachi, Nara, 630-8506,
Japan; e-mail: scarab@cc.nara-wu.ac.jp

1

Environmental Entomology and Zoology, Graduate School of Life and Sciences, Osaka Prefecture University, Sakai, Osaka,599-8531, Japan; e-mail: UHI06205@nifty.com

2

Bird Rescue Society of Japan, 3918 Oyamadacho, Kawachinagano, Osaka, 586-0094, Japan; e-mails: alba@bird-rescue.jp,
h-matsumuro_s5824@ab.auone-net.jp

3

4

Rikkyo University, 3-34-1 Nishi-Ikebukuro, Toshima-ku, Tokyo, 171-8501, Japan; e-mail: keisuke@rikkyo.ac.jp

Key words. Lepidoptera, Tineinae, Monopis, Niditinea, niche, larviparity, bird nests, Ural owl, Japan
Abstract. Some moth larvae feed not on plants but on keratin and/or chitin produced by animals. These substances are polymers
and are commonly found in bird nests as feathers and raptor pellets. Many qualitative studies have examined the association of
keratin/chitin feeding moths with bird nests. However, few studies have quantified the species composition with respect to type ofnest and habitat. Hence, we have studied the degree to which the niches of these moths differ in terms of type of nest and habitat.
We set-up open-top nest boxes for the Ural owl Strix uralensis (damp exposed nests from which owl chicks were fledged successfully) and artificial bird nests (mesh bags filled with duck down to imitate dry exposed nests) in a deciduous forest and artificial
bird nests in an urban setting in Aichi Prefecture, central Japan, and collected the contents of the nests every two months from
June to December 2014. We recorded 592 individuals of five keratin/chitin feeding moth species (Tineidae, Tineinae) from thecontents. Using non-metric multidimensional scaling and cluster analysis of the relative abundances of individual species in eachtype of nest in forest and urban settings, these species were classified into three groups: (1) Monopis longella and M. congestella
inhabiting forest, (2) M. flavidorsalis and Niditinea tugurialis mainly in damp exposed nests in forest and (3) N. piercella mainly indry exposed nests in urban areas. This classification was compared with findings of other studies. As a result, these moths probably have different niches with respect to nest type (damp or dry), keratin/chitin as a source of food (raptor pellets or feathers),
and habitat (forest or urban area). Furthermore, we suggest that the evolution of larviparity in M. congestella might be related toits preference for feathers as a source of food for the development of its larvae.

INTRODUCTION

Bird nests harbour highly diverse insects including
Hemiptera, Diptera, Siphonaptera, Coleoptera, Hymenoptera and Lepidoptera (Nordberg, 1936; Woodroffe, 1953;
Hicks, 1959, 1962, 1971; Krišrosík et al., 1994). Of these
insect orders, the relationship of Lepidoptera (moths) withbird nests has been comparatively well studied because
moths tend to be noticeable and abundant in nests (Jalava,
1980; Jaworski et al., 2012; Nasu et al., 2012a, c; Boyes& Lewis, 2019). Moth larvae in nests are classified mainly
into plant- and animal-material feeders (Robinson, 2004;
Nasu et al., 2012a, c). The former feed on dead leaves,grass, fibres, moss and liverwort (albeit not plants), while
the latter feed on faeces, raptor pellets, hair and feathers,
which contain keratin and chitin. Hence, it seems likely thatthe moth fauna in bird nests depends on the composition of
the nest contents (Nasu et al., 2012a, c). However, Boyes& Lewis’ (2019) recent comparison of moth assemblages

in concealed nests of five passerine species in mature oak
woodland in mid-Wales, UK, revealed no evidence of theoccurrence and abundance of species of moths in nests depending on nest contents. They offer two possible explanations. First, nest discovery by ovipositing female moths
may be largely a matter of chance. Second, detritovores
may be unspecialized feeders. This study focused on keratin/chitin feeding moths (animal-material feeders) in orderto clarify the association of moths with bird nests.
According to Robinson (2004), keratin/chitin feeding
moths belong to the subfamilies Tineinae and Hieroxestinae of the family Tineidae. Most species of Tineinae are
keratin and/or chitin feeders, while only a few of the Hieroxestinae feed on chitin. Of the keratin and/or chitin feeding species of Tineinae, Monopis, Niditinea and Tinea arechiefly associated with bird nests. Robinson (2004) also
shows that keratin/chitin feeding moths are likely to occupy different niches with respect to nest type, i.e., damp

Final formatted article © Institute of Entomology, Biology Centre, Czech Academy of Sciences, České Budějovice.
An Open Access article distributed under the Creative Commons (CC-BY) license (http://creativecommons.org/licenses/by/4.0/).

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concealed, dry concealed, damp exposed, or dry exposed;
concealed nests are those in holes in trees (including nest
boxes), while exposed nests are constructed in trees andhedges. For example, the Monopis congestella-group and
M. monachella-group are found in damp exposed and dry
exposed nests, respectively, while the Niditinea fuscella-group are found in nests of both types. The Niditinea
piercella-group occur only in concealed nests. However,Robinson (2004) does not provide quantitative data supporting these conclusions.
Since Robinson (2004), much information has been collected about keratin/chitin feeding moths in bird nests,
especially in Japan (e.g., Nasu et al., 2007a, b, 2008a, b,
2012a, b, c). In these studies moths were reared on theorganic contents of concealed and exposed nests of 16
species of birds in forests and urban areas and Monopis
longella (Walker) [= pavlovskii (Zagulyaev)], M. congestella (Walker), M. flavidorsalis (Matsumura), Niditinea
tugurialis Meyrick [= baryspilas (Meyrick)], N. striolella(Matsumura), Tinea translucens Meyrick and Tineola bisselliella (Hummel) fed on the keratin/chitin in the nests ofthese birds. Nasu (2012) summarizes the associations ofthese moths with bird nests in terms of type of nest andhabitat as follows: M. longella and M. congestella occur inconcealed and exposed nests in forests, while M. flavidorsalis and N. striolella are found only in concealed nests inforests; in contrast, Tinea translucens and Tineola bisselliella only occur in concealed and exposed nests in urban
areas. However, the studies by Nasu and colleagues did notquantitatively or systematically assess the degree to whichthese moths depend on concealed and exposed nests. Furthermore, the specimens identified as N. tugurialis probably included N. piercella Bentinck, so information on theoccurrence of this species in bird nests is unreliable (Nasu& Sato, 2016).
Larviparity, or laying fully developed first-instar larvae instead of eggs, is very rare in Lepidoptera (Hinton,
1981; Heppner, 2009). However, it is relatively common
in Monopis, a keratin/chitin feeding genus with 90 species
worldwide (Robinson, 2019), 12 of which are known to belarviparous (Xiao & Li, 2006). Three species of Monopis
are recorded in Japan (see above; Sakai, 2013), and one,
M. congestella, is larviparous (Robinson, 1988, 2004;
Nasu et al., 2008a). Robinson (2004) has hypothesized thatlarviparity in Monopis evolved due to (1) high mortality ofeggs and neonates due to intense predation by mites in bird
nests, and (2) high cost of finding more than one oviposition site, or bird’s nest. If there is a difference in nests exploited by larviparous and oviparous Monopis moths, thenthere could be another reason.
This study determines the degree to which keratin/chitin
feeding moths have different niches in terms of the typesof nests or habitats in which they are constructed. This involved sampling moths quantitatively and systematicallyin a deciduous forest and an urban area using nest boxes
and artificial bird nests. The open-top nest boxes were designed for the Ural owl Strix uralensis Pallas, while theartificial bird nests were made from duck-down stuffed

doi: 10.14411/eje.2019.045

into a plastic mesh bag (Robinson & Nielsen, 1993). The
owl nest boxes and artificial bird nests were categorized
as damp exposed type and dry exposed type, respectively,
following Robinson’s (2004) classification. We compared
the composition of keratin/chitin feeding species of mothin the two types of nests in different habitats.
MATERIALS AND METHODS
Ecology of the Ural owl

Paired Ural owls usually nest in cavities in tree trunks. Females
lay two to four eggs in March to April and incubate them for
27 to 32 days (Nakamura & Nakamura, 1995). Prey for chicks
are small mammals, birds, frogs, newts, lizards and centipedes(our unpublished data). Chicks leave the nest 30 to 34 days after
hatching.
Collection of the organic contents of owl nest boxes

Nine open-top plastic boxes (26 cm wide, 26 cm long, 30 cmdeep) were installed in trees 2 to 3 m above ground level in asecondary deciduous forest (80 to 360 m a.s.l.) extending across
Toyota and Okazaki Cities, Aichi Prefecture, central Japan in
May 2013 (Fig. 1). Several twigs were put in each box as nest
material. In spring 2014, Ural owls bred in each box, with one or
two chicks fledged by late May. All or half the organic contents(feathers, owl pellets, faeces and other detritus) were collectedfrom each of three or four randomly chosen boxes on 7 June, 17
August, 18 October and 6 December 2014; as a result, the organic
contents in five boxes were collected twice (Table 1). Feathers
were present in only two boxes in June and August. The quantityof nest material collected at any one time ranged from 0.6 to 3.1
litres.
Sampling using artificial birds’ nests

Artificial birds’ nests were made according Robinson & Nielsen (1993) using plastic mesh bags (16 × 20 cm in size; 3 × 3 mmin mesh size) stuffed with 10 g of dry duck down bound together
with 5 g of beer yeast and some water. Four artificial nests werehung on tree branches (several hundred meters apart; 90 to 120 ma.s.l.) at 1.5 m above the ground in a secondary deciduous forest
in the Toyota City Nature Sanctuary, Aichi Prefecture on 7 June,
17 August and 18 October 2014 (hereafter referred to as forest
artificial nests; Fig. 1). To compare the species composition of
the moths in the forest and that in the urban area, three artificial
bird nests were set-up at the Toyota Yahagi River Institute in theurban area of Toyota City on 2 June, 18 August and 16 October
2014 (hereafter referred to as urban artificial nests; Fig. 1); thissite is 400 m from the nearest green park. Two artificial nests
were hung on the branches of trees at this site, while the third
was hung under the eaves of a building. The nests were retrieved
approximately two months later on 17 August, 18 October or 6
December 2014.
Rearing

Organic contents collected from each owl nest box on each occasion they were sampled were put in one to three plastic boxes(16 cm wide, 22 cm long, 8 cm deep) according to the amount.
To ventilate the boxes, the centre of the lid of each box was cut
away to form 4-cm square window, which was covered by a sheetof fine plastic mesh. Duck down from each artificial nest was also
kept in a plastic box of the same type. These boxes were stored
under semi-natural temperature and light-dark photoperiod conditions (adjusted to monthly average values in field) in an incubator
for two to seven months according to the collection date; materials collected in June, August, October and December were kept

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doi: 10.14411/eje.2019.045

Fig. 1. Location of owl nest boxes (solid circles) and artificial bird nests (open circles). Four and three artificial nests (mesh bags stuffedwith duck down) were located in a deciduous forest in the Toyota City Nature Sanctuary (forest artificial nests) and at the Toyota YahagiRiver Institute in an urban area (urban artificial nests), respectively.

for two, two, seven and five months, respectively. Adult moths
emerging during these periods were identified to species and thenumbers were counted. The number of cocoons of M. congestella
within which a dead or alive mature larva or a pupa existed was
also counted. Niditinea tugurialis and N. piercella were distinguished by their genitalia according to Nasu & Sato (2016).
Niditinea piercella is sometimes treated as a junior synonym of
N. striolella (Matsumura) (Karsholt & Razowski, 1996; Gaedike,
2000, 2019). However, the drawings of male and female genitalia
identified as those of N. pacifella (which is a junior synonym of
N. striolella) in Zagulyaev (1960) differ greatly from those of N.
piercella in Pelham-Clinton (1985). According to these authors,
N. striolella occurs in the Russian Far East and Japan, while N.
piercella is found in Britain and continental Europe. Accordingly,
Robinson (2009) recommends keeping the two species separate
until material from across the continent can be compared. Recently, Sakai (2013) and Nasu & Sato (2016), based on the structureof the male and female genitalia of Japanese specimens of possible N. striolella and N. piercella, conclude that these two species
are distinctly different. Therefore, this paper treats N. piercellaand N. striolella as two species.

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Table 1. Details of owl-nest contents and when collected. Contentsof nests 1, 2, 4, 6 and 7 (bold-face) were collected twice on different dates. Half the contents of these nests was sampled the firsttime and the remaining contents the second time. Nest numbers
correspond to those in Fig. 1.
Collection date

Nest No.

7 June 2014

1
2
3
4
1
5
6
7
4
7
8
2
6
9

17 August 201418 October 2014

6 December 2014

Nest content collected
Amount (l)Proportion
0.6
1.6
4.8
1.8
0.6
2.7
0.5
2.5
1.6
2.5
3.1
1.6
0.5
3

1/2
1/2
1
1/2
1
1
1/2
1/2
1
1
1
1
1
1

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Fig. 2. Relative abundance of moths of Tineinae reared from thecontents of owl nest boxes, forest artificial nests and urban artificial nests. Numbers in parentheses represent the total numberof moths. Nine owl nest boxes were located in a forest covering
a geographic region, while forest artificial nests and three urban
artificial nests were located in a small forest and at an institute inan urban area, respectively (see Fig. 1).

Statistical analysis

To determine whether there are differences in the niches of thedifferent species of moths we carried out a non-metric multidimensional scaling (NMDS) and cluster analysis using arcsinetransformed proportions of individual species in owl nest boxes,
forest artificial nests and urban artificial nests. Both statisticalprocedures were based on a Bray-Curtis dissimilarity matrix.
Niche differences were visualized using a NMDS ordination plotin 2D space. Moth species with similar niches were grouped bycluster analysis using Ward’s linkage method. These procedures
were performed using IBM SPSS ver. 25 (IBM 2017).

RESULTSA total of 592 individuals of five species of Tineinae, i.e.,
M. longella, M. congestella, M. flavidorsalis, N. tugurialis
and N. piercella, were obtained from owl nest boxes andartificial nests (Fig. 2). The composition of species in the
owl nest boxes, forest artificial nests and urban artificial
nests differed. The dominant species in owl nest boxes was
M. longella (78.5%), followed by N. piercella (11.0%). The
dominant species in forest artificial nests was also M. longella (75.4%), followed by M. congestella (24.1%), which
was not found in owl nest boxes. Monopis flavidorsalis wasrecorded only from owl nest boxes (6.3%). Most moths inurban artificial nests were N. piercella (99.5%) with theother species M. longella (0.5%). Niditinea piercella wasalso common in owl nest boxes (11.0%), but not in forest
artificial nests. Niditinea tugurialis was found in both owl
nest boxes (4.2%) and forest artificial nests (0.5%), but notin urban artificial nests.
The NMDS ordination plot in 2D space (Fig. 3A) had
a stress value of 0.0091. This near-to-zero value indicatesthat the plot is a reliable representation (Clarke, 1993).
The diagram appears to show that the five species of Tineinae are classified into three groups. Monopis longellaand M. congestella belong to the same group, while M. flavidorsalis and N. tugurialis are classed together. Niditinea
piercella forms a separate group. This classification was
confirmed by cluster analysis (Fig. 3B). The dendrogram

Fig. 3. NMDS ordination plot (A) and dendrogram obtained using
cluster analysis (B) of the relative abundances of individual species in each nest (Fig. 2). The five species are classified into threegroups: (1) Monopis longella and M. congestella, (2) M. flavidorsalis and Niditinea tugurialis, and (3) N. piercella. The vertical dottedline on panel B indicates the level of 50% dissimilarity.

and the NMDS diagram both put the five species into threegroups.
No adult moths emerged from the contents collectedfrom owl nest boxes or urban artificial nests in December.
Monopis longella adults were reared from owl nest contents collected in June, August and October (Fig. 4A), andfrom forest artificial nests in August, October and December (Fig. 4B), which indicates two peaks in occurrence.
Monopis congestella and M. flavidorsalis were recorded
in forest artificial nests in August, October and December,and in owl nest boxes in June, August and October. Larvaeof M. congestella that were collected in December grew
into final stage and used silk to construct a cocoon of down,
but all of them subsequently died from unknown causes. In
comparison to the species of Monopis, N. piercella and N.
tugurialis were present for short periods from June to August and June to October, respectively.
DISCUSSION

We found five species of Tineinae (Tineidae) that feed on
keratin/chitin in the organic contents of owl nest boxes andartificial nests (i.e., duck down stuffed in mesh bags) placedin a forest and urban areas. These moths were assigned to
one of three groups, which include: (1) M. longella and M.
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Fig. 4. Bimonthly composition of the species recorded in owl nest boxes, forest artificial nests and urban artificial nests. Abundances ofmoths are expressed in terms of the number of individuals per litre of the organic content of a nest or per artificial nest.

congestella, (2) M. flavidorsalis and N. tugurialis, and (3)N. piercella (Fig. 3). However, unlike M. longella, most M.
congestella were found in forest artificial nests, which is
not reflected in the above three-group classification.
Robinson (2004) suggests that the type of nest and foodresource are important in determining niche separationin keratin/chitin feeding moths and defines four types ofnests: damp concealed, dry concealed, damp exposed anddry exposed. Based on his definitions, owl nest boxes in
the present study are classed as damp exposed, because the
top is open and drainage is poor, so the organic contents inthe box are wet. Hence, these nest boxes are unlike natural
owl nests, which are usually dry and concealed in cavities
in tree trunks. In contrast, mesh bags filled with down toimitate bird nests are classed as dry exposed, because thebags were hung in trees or in the eaves of a building, so thedown remained dry although left outdoors for two months.
Robinson (2004) describes the food sources of keratin/chitin feeding larvae as either compact or loose. The former
includes carnivore droppings, raptor pellets and guano,
while the latter mainly comprises hair, fur and feathers. In
this study, the contents of owl nest boxes are classed as acompact resource, while those in the artificial bird nests areclassed as a loose resource.
Based on Robinson’s (2004) definitions together with
habitats (forest and urban) three groups of Tineinae moths
may be characterized as follows: (1) M. longella and M.
congestella inhabiting forests, (2) M. flavidorsalis and N.
tugurialis exploiting damp exposed nests containing compact sources of food in forest, and (3) N. piercella mainly using dry exposed nests containing loose resources inurban areas. There may be doubts about the differences inniches of these moths, because the forest and urban artificial nests were in one forest and one urban area, respectively, whereas the owl nest boxes were spread across ageographical region (Fig. 1). Nevertheless, the urban site
is regarded as typical because it is 400 m from the nearest green park and little affected by semi-natural environments. Furthermore, the forest where the owl nest boxes
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were deployed is located in a wide stretch of hills coveredby secondary deciduous forest dominated by oak, Quercus
serrata. Hence, even if artificial nests had been set at othersites in the hills as the owl nest boxes were, some species
might have been added, but the dominant species might
have been little changed. In fact, N. striolella is the only
Tineinae species recorded from owl nests in Japan, but was
not recorded in the present study; in addition, this species
is recorded only infrequently (Nasu et al., 2012b). Consequently, the differences in the niches of the five species ofTineinae seem acceptable, albeit insufficiently supported.
Therefore, we compare the nature of the niches of thesemoths reported here with that reported in other studies.
In Japan, Nasu et al. (2007a, 2008b, 2012a, c) report thatmany more M. longella than M. congestella emerge fromthe organic contents of Ural owl nests (and concealed andopen nest boxes) and of the nest boxes of the varied tit (Sittiparus varius Temminck & Schlegel). More M. longellathan M. congestella are usually recorded from nest boxesof the Japanese tit (Parus minor Temminck & Schlegel,mis-described as P. major L.) and nests of Styan’s grasshopper-warbler (Locustella pleskei Taczanowski) (Nasuet al., 2007b, 2012c). Sittiparus varius and P. minor nest
boxes are dry and concealed, whereas those of L. pleskei
are dry and exposed. These passerines are insectivorous
and their nests contain faeces, fur and feathers. All thesenests were located in secondary forests. In Korea, Lee etal. (2016) reared as many M. congestella as M. longellafrom the contents of mesh bags containing feathers, pellets and faeces of a raptor in a forest. In addition, Nasu etal. (2007b, 2012b; Y. Nasu & S. Murahama, unpubl. data)
reared M. longella, but not M. congestella, from pelletsof the common buzzard (Buteo buteo (L.)), sparrowhawk(Accipiter nisus (L.)) and common kestrel (Falco tinnunculus L.) lying on the ground, which indicates that M. longella prefers more compact sources of keratin/chitin thanM. congestella. Combining these and our results, we conclude that (1) both M. longella and M. congestella are forest inhabitants, (2) M. longella has a larger niche breadth,Sato et al., Eur. J. Entomol. 116: 442–449, 2019

exploiting damp and dry concealed nests and dry exposed
nests and consuming both compact and loose sources offood and (3) M. congestella has a smaller niche breadth,
exploiting dry concealed and exposed nests and loose
sources of food. However, this conclusion is partly inconsistent with Robinson’s (2004) classification as he considers M. longella to be a damp-exposed nest exploiter. This
inconsistency is possibly due to small sample size.
Monopis flavidorsalis and N. tugurialis [= baryspilas] arerecorded from the pellets of common buzzards and sparrow hawk lying on the ground (Nasu et al., 2007b, 2012b).
Taking our results into consideration, these two species arelikely to prefer compact sources of food regardless of nest
type. In addition, N. tugurialis is not only recorded from
Ural owl nests (Nasu et al., 2007a, b, 2012a) but also thedry concealed nests of brown hawk-owl (Ninox scutulata (Raffles)) (Nasu et al., 2012a), Japanese tit nest boxes(Nasu et al., 2008b; Nasu & Saito 2008), varied tit nest
boxes (Nasu et al., 2008b), damp exposed nests of the Oriental stork (Ciconia boyciana Swinhoe) (Nasu et al., 2010)
and dry concealed nest boxes of the Oriental dollar bird
(Eurystomus orientalis L.) (Nasu et al., 2012c). Moreover,
adults and larvae of N. tugurialis are recorded from guanoof the greater horseshoe bat (Rhinolophus ferrumequinum(Schreber)) (Nasu et al., 2016). These records might indicate that this species uses a variety of types of nest. However, specimens identified as N. tugurialis in studies before
2012 probably include N. piercella, because prior to thatdate genitalia were not used to distinguish between thesemorphologically very similar species (Nasu & Sato, 2016).
This prevents a more rigorous assessment of the types ofnests N. tugurialis tends to exploit. According to Robinson (2004), N. tugurialis exploits both damp-exposed anddry-exposed nests. If so, the niches of N. tugurialis and M.
flavidorsalis might be different.
Our results show that N. piercella has a very different
niche from the remaining four species, depending primarily on dry exposed nests containing loose sources offood in urban areas. Nevertheless, it is puzzling that noN. piercella adults emerged from forest artificial nests despite the fact that this species inhabits forests and exploits
owl nest boxes (Fig. 2). This might be due to the artificial
nests only sampling a small area of forest. Contrary to our
results, Robinson (2004, 2009) considers N. piercella tobe an exploiter of concealed-nests based on his and otherstudies (Jalava, 1980; Pelham-Clinton, 1985; Robinson &Nielsen, 1993; Robinson, 2004). Results of recent studiesof tawny owl (Strix aluco L.) nests in Poland (Jaworski etal., 2012) and concealed nests of passerines in mid-Wales,
UK (Boyes & Lewis, 2019) support his proposal. However, specimens identified as N. striolella are probably N.
piercella as mentioned above. In Japan, N. striolella isneither frequent nor dominant in owl nests (Nasu et al.,2012a), but is recorded occasionally in Oriental dollar
bird nests (Nasu et al., 2012c). Another problem is that N.
piercella was once confused with N. tugurialis in Japan as
mentioned above (Nasu & Sato, 2016). To properly define

doi: 10.14411/eje.2019.045

the niche of N. piercella we must collect further data andidentify specimens more carefully.
Based on larval occurrence over more than three months,
the five species of Tineinae in the present study are likely
to be multivoltine. For M. longella, the two peaks in theyear in larval occurrence (Fig. 4) probably indicate twogenerations. Monopis longella and M. congestella are believed to overwinter in the larval stage, because the larvae recorded in owl nest boxes and artificial bird nests in
October and/or December developed into mature larvae or
adults by May of the next year under semi-natural conditions. Overwintering of M. longella, M. congestella andM. flavidorsalis in the larval stage is reported by Nasu etal. (2007b, 2012b). Nevertheless, no adults emerged fromorganic contents collected from owl nest boxes in December (Fig. 4). One possible reason is that keratin/chitin resources had been mostly consumed or decomposed by thattime. If so, the result might indicate that there is intensecompetition for food among larvae in owl nests. Although
we did not record the overwintering stage of N. piercellaand N. tugurialis, other studies report overwintering in thelarval stage (Nasu et al., 2012c; Sakai, 2013).
Monopis congestella is known to be larviparous (Robinson, 1988, 2004; Nasu et al., 2008a). Females of M. congestella lay fully-developed first instar larvae, instead oflaying eggs. Although larviparity is rare in Lepidoptera(Hinton, 1981; Heppner, 2009), it often occurs in Monopis;
12 of 90 species of Monopis are larviparous (Xiao & Li,
2006). Robinson (2004) presents two hypotheses for theevolution of larviparity in Monopis: (1) high mortality ofeggs and newly-hatched larvae in bird nests due to predation by mites, and (2) females have difficulty finding morethan one oviposition site or bird’s nest. However, thesehypotheses do not explain why the majority of Monopis
species are oviparous. In M. congestella, the evolution oflarviparity might be related to the main food source being
feathers. The quantity of feathers in bird nests is usuallysmall, so larvae that can pre-empt an eventual shortage offood have an advantage. To corroborate this hypothesis,
we need to compare the feeding habits of larviparous andoviparous Monopis.
In conclusion, this study indicates that Tineinae moths
associated with bird nests differ in their niches in termsof type of nest (concealed or exposed), source of keratin/
chitin (raptor pellets or feathers) and habitat (forest or
urban area). However, some problems remain unresolved.
First, we cannot explain why N. piercella did not exploit
the exposed nests in a forest where it lives even though it
exploits such nests. Second, the degree to which Tineinae
larvae compete for sources of keratin/chitin in owl nests is
unclear. Third, the factors contributing to the evolution oflarviparity in Monopis are unclear. We will address these
problems in the near future.
ACKNOWLEDGEMENTS. We thank T. Mano for setting up andretrieving artificial bird nests at the Toyota Yahagi River Instituteand T. Sugiyama for allowing us to collect the organic contentsfrom owl nest boxes. Toyota City Nature Sanctuary permitted us
to collect moths in the forest. The Research Institute of the Envi-

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Sato et al., Eur. J. Entomol. 116: 442–449, 2019ronment, Agriculture and Fisheries, Osaka Prefecture kindly provided us with duck down. M. Sakai answered our questions about
Niditinea taxonomy. This work was supported financially by aGrant-in-Aid for Scientific Research (KAKENHI) from the Japan
Society for the Promotion of Science (No. 25291090).

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Received April 30, 2019; revised and accepted November 11, 2019
Published online November 27, 2019449