Relationships Between Nest-Dwelling Lepidoptera and Their Owl Hosts

Ornithol Sci 11: 77 – 85 (2012)

ORIGINAL ARTICLE Relationships between nest-dwelling Lepidoptera and their owl hosts

Yoshitsugu NASU1,#, Shiro MURAHAMA2, Hiroyuki MATSUMURO3, Keisuke UEDA4, Toshiya HIROWATARI5 and Yutaka YOSHIYASU6

1 153–2, Nakado, Hashimoto, Wakayama 648–0023 Japan 2 Wildlife Conservation Laboratory Co. Ltd., 1–2–40–402, Miyahara, Yodogawa, Osaka 532–0003 Japan 3 Japan Bird Rescue Association, 3918, Oyamada-cho, Kawachinagano, Osaka 586–0094 Japan 4 College of Science, Rikkyo University, 3–34–1, Nishi-Ikebukuro, Toshima, Tokyo 171–8501 Japan 5 Entomological Laboratory, Osaka Prefecture University, 1–1, Gakuen-cho, Naka-ku, Sakai, Osaka 599–8531 Japan 6 Laboratory of Applied Entomology, Kyoto Prefectural University, Shimogamo, Kyoto 606–8522 Japan

Abstract Lepidoptera fauna of five owl species nests were investigated in Japan. ORNITHOLOGICAL Seventeen moth species were identified: Niditinea striolella (Tineidae), Agonopterix SCIENCE sp. (Elachistidae) from Blakiston’s Fish Owl, Ketupa blakistoni, nests; Monopis lon- © The Ornithological Society gella (= pavlovskii), M. flavidorsalis, M. sp., M. congestella, Niditinea baryspilas, N. of Japan 2012 striolella, N. sp. (Tineidae), Martyringa ussuriella (Oecophoridae), Mabra charonialis (Crambidae), Pyralis regalis (Pyralidae) from Ural Owl, Strix uralensis, nests; Tinea translucens, Niditinea baryspilas (Tineidae) from Brown Hawk-Owl, Ninox scutulata, nests; Opogona sacchari, O. sp., Phaeoses sp. (Tineidae) from Collared Scops Owl, Otus lempiji, nests; and Opogona sacchari, Phaeoses sp., Setomorpha sp. (Tineidae), Endotricha theonalis (Pyralidae) from Ryukyu Scops Owl, Otus elegans, nests. The moth nest fauna varied among owl species. The differences were related to owl prey (fish, small animals and birds, insects) and habitats (urban area, forest), and the tineid species selecting the nest. Tineids are presumed to decompose keratin found in owl nests and help maintain the cleanliness of the nest chamber, and such relationships between tineids and owls may be mutualistic. Rapid burrowing into owl nest materials by tineids may reflect a strategy to avoid being preyed upon by the nest owners.

Key words Detritus, Ecosystem engineer, Nest-dwelling, Symbiosis, Tineidae

Birds modify their environments by activities such alba (Hicks 1959), and five species of Tineidae have as nest construction, and hence act as ecosystem been found in the nests of Finnish Ural Owl, Strix engineers (Jones et al. 1994). Other organisms, espe- uralensis (Jalava 1980). cially insects, utilize bird nests as habitats (Nordberg From 2006 to 2011, the lepidopteran fauna in the 1936; Woodroffe 1953; Hicks 1959, 1962; Whelan et nests of five owl species: Blakiston’s Fish Owl, Ket- al. 2008); however, few studies have focused on the upa blakistoni, Ural Owl, Brown Hawk-Owl, Ninox relationships between insects and birds. scutulata, Collared Scops Owl, Otus lempiji, and Only fragmentary investigations have been made Ryukyu Scops Owl, O. elegans, was surveyed in of Lepidoptera in owl nests; nevertheless, Tinea sp. Japan. As a result of the investigations, 17 moth spe- (Tineidae), Gerdana sp. (Blastobasidae) and Alucita cies were identified (12 tineids, one oecophorid, one huebneri (Alucitidae) have been reported from nests elachistid, two pyralids and one crambid) of which, of North American Long-eared Owl, Asio otus, Hof- five tineids and one oecophorid had been reported mannophila pseudopretella (Oecophoridae) has been previously from Ural Owl nests by Nasu et al. (2007a, noted from the nests of European Barn Owl, Tyto 2008) and Huang et al. (2011). Here we provide data on the 17 moth species found, and discuss the role of (Received 26 March 2012; Accepted 6 May 2012) owls as ecosystem engineers and the relationships # Corresponding author, E-mail: [email protected] between the moths and the birds.

77 Y. NASU et al.

Fig. 1. Map of owl nest localities, Japan.

MATERIALS AND METHODS All of the moth specimens are preserved in the collection of the Entomological laboratory of Osaka Detritus was collected from the bottom of a range Prefecture University, Sakai, Osaka, Japan. of natural nests, and detritus and wood chip remains (provided as artificial nest material) was collected RESULTS from nest boxes from which young owls fledged (see Fig. 1). The detritus was put into rearing cases at 1) Blakiston’s Fish Owl 25°C in an incubator or at room temperature with a Two nests supported the tineid moth Niditinea stri- natural photoperiod cycle, the larvae in the detritus olella (Fig. 2g), and similarly two nests supported the were allowed to develop and the adult moths that elachistid moth Agonopterix sp. (Fig. 2n; Table 1). In emerged were then identified. addition, two tineid larvae, probably N. striolella, Most owl nests studied were in forests, although were found in one nest. The nest detritus consisted Brown Hawk-Owl nests were in urban areas. The mainly of decaying wood chips, a few owl down nest numbers and dates were as follows: Blakiston’s feathers, and fish scales. Fish Owl: in 2007–2011, one natural nest and 13 nest boxes were investigated in Hokkaido (for con- 2) Ural Owl servation reasons exact nest localities are not given Moths were incubated from most nests, with a this endangered species); Ural Owl: in 2006–2011, range of species represented: seven tineids, four two natural nests and 57 nest boxes in 21 localities, Monopis including an unknown species (Fig. 2c), 11 prefectures; Brown Hawk-Owl: in 2009 and 2010, three Niditinea including an unknown species (Fig. five natural nests and one nest box in six localities, 2i), one oecophorid, Martyringa ussuriella (Fig. 2m), three prefectures; Collared Scops Owl: in 2007 and one crambid, Mabra charonialis (Fig. 2o) and one 2008, nine nest boxes in Kunigami-son, Okinawa Is., pyralid, Pyralis regalis (Fig. 2p; Table 2). Of these, Okinawa Prefecture; Ryukyu Scops Owl: in 2007 and Monopis longella (= pavlovskii) (Fig. 2a), M. flavi- 2008, six nest boxes in Kunigami-son, Okinawa Is., dorsalis (Fig. 2b), M. congestella (Fig. 2d), Niditinea Okinawa Prefecture, and in 2007–2009, nine natural baryspilas (Fig. 2f) and N. striolella predominated. A nests and two nest boxes on Minami-daito Is., great number of Monopis longella were reared, with Okinawa Prefecture. over 328 moths per nest. From the nest detritus col-

78 Nest-dwelling Lepidoptera and owls

Fig. 2. Lepidoptera from owl nests (a-l, Tineidae; m, Oecophoridae; n, Elachistidae; o, Crambidae; p, q, Pyralidae). a, Monopis longella (= pavlovskii); b, M. flavidorsalis; c, M. sp.; d, M. congestella; e, Tinea translucens; f, Niditinea baryspilas; g, N. striolella; h, Opogona sacchari; i, Niditinea sp.; j, Opogona sp.; k, Phaeoses sp.; l, Setomorpha sp.; m, Martyringa ussuriella; n, Agonopterix sp.; o, Mabra charonialis; p, Pyralis regalis; q, Endotricha theonalis.

Table 1. Number of Lepidoptera from Blakiston’s Fish Owl, Ketupa blakistoni, nests in Japan.

Date detritus Tineidae Elachistidae Locality Nest collected Niditinea striolella larval case Agonopterix sp. Hokkaido Nest box Oct. 26, 2007 6* 2*** Oct. 26, 2007 10* 1*** Aug. 11, 2011 2**

* larvae, moths developed in Feb., 2008; ** probably Niditinea striolella; *** adults.

79 Y. NASU et al. 2 Mabra charonialis 2 regalis Pyralis Pyralidae Crambidae 1 1+ 1+ 1+

ussuriella Martyringa Oecophoridae

1 sp. Niditinea 11 5 striolella Niditinea 5+ 1 1+ 5+ 1+ 5 1+ 1+ 100+ 23 66 30+ 23 1 2 58 5 3 5 3 Niditinea baryspilas 6+ 9+ 59 58+ 6 57+ 18

2 18 22 9 8 40 Monopis congestella Tineidae 3 2 1+ 1 sp.

Monopis 6+ 1+ 1+ 80 12 38+ 6

2 1 5 1 1 9 Monopis flavidorsalis 11 + 1 115 + 7+ 37 328+ 28+ 7+ 18 13+ 1 9+ 25+ 64+ 2 26

8+ 40+ 6 2 14 3 166 43 71 36 24 8 8 100+ 1 2+ 13 ) pavlovskii Petaurista leucogenys (= Monopis longella Strix uralensis , nests in Japan. *** * ** nest reused by collected *** Date detritus Date detritus Jul. 9, 2007 May 8, 2007 Jun. 26, 2007 Jul. 12, 2009 Jul. 12, 2009 Jul. 17, 2010 Jul. 17, 2010 Jul. 7, 2007 Jul. 4, 2009 Jun. 16, 2007 Aug. 8, 2010 Jun. 16, 2007 Jun. 16, 2007 Jul. 6, 2008 Jul. 18, 2009 Aug. 8, 2010 Jul. 6, 2008 Jun. 16, 2007 Aug. 7, 2010 Aug. 7, 2010 Jul. 9, 2011 Aug. 26, 2007 Aug. 26, 2007 Jul. 9, 2011 Parus spp.; Nest Natural nest Nest box Jul. 20, 2009 Nest box Jun. 26, 2007 Nest box Jul. 17, 2010 Nest box Jul. 12, 2009 Nest box Jul. 12, 2009 Nest box Jul. 12, 2009 Nest box Jun. 25, 2006 Nest box Jul. 11, 2010 Nest box Jun. 25, 2006 Natural nest Nest box Jun. 6, 2008 nest reused by ** Toyota Nishio Kira Kouda Takashima Nest box Sep. 29, 2006 Ujitabaru Nest box Jul. 12, 2009 Hirakata Ikoma Shijonawate Nest box Jul. 6, 2008 Katano Daito Kawachinagano Nest box Jul. 19, 2009 Miyakonojo Nest box Aug. 1, 2010 Number of Lepidoptera from Ural Owl, Locality AomoriYamanashi Aomori Fujiyoshida Nest box May 2, 2007 Nagano Nagano Aichi Okazaki Shiga Kyoto Osaka Minoh Nara Wakayama Hashimoto Nest box Jun. 17, 2006 Miyazaki nest detritus in 2006; Table 2. Table *

80 Nest-dwelling Lepidoptera and owls

Table 3. Number of Lepidoptera from Brown Hawk-Owl, Ninox scutulata, nests in Japan.

Tineidae Date detritus Locality Nest collected Tinea Niditinea translucens baryspilas Kyoto Kyoto Nest box Oct. 15, 2010 1 Miyazaki Miyakonojo Natural nest Aug. 1, 2009 10+* Miyazaki Aug. 3, 2009 2 22+

* larvae

Table 4. Number of Lepidotpera from scops owls, Otus spp., nests in Okinawa, Japan.

Tineidae Pyralidae Date detritus Owls Locality Nest collected Opogona Opogona Phaeoses Setomor- Endotricha sacchari sp. sp. pha sp. theonalis

Collared Kunigami, Nest box Jul. 8, 2007 1 Scops Owl, Okinawa Is. Jul. 8, 2007 3 1 Otus lempiji Jul. 19, 2007 1 4 Jul. 19, 2007 7+ Jul. 13, 2008 1

Ryukyu Kunigami, Nest box Jul. 15, 2008 2 2 Scops Owl, Okinawa Is. Otus elegans Minami-daito Is. Natural nest Jun. 24, 2007 1 Jun. 24, 2007 3 Jun. 24, 2007 1+ Jun. 24, 2007 1+ Jun. 24, 2007 5 Jun. 27, 2008 1

Nest box Jun. 24, 2007 9+ lected in Yamanashi Prefecture in May 2007 alone, pyralid, Endotricha theonalis (Fig. 2q), was also over 115 M. longella were reared. The detritus from reared. The nest detritus consisted of decaying wood these nests consisted of decaying wood chips, feath- chips, down, pellets, food remains (crushed insects) ers, down, pellets, food remains (birds, small ani- and droppings. mals) and droppings. In all owl nests, the tineid adults reared from the detritus flew poorly, fell on disturbance, crash-dived 3) Brown Hawk-Owl and burrowed into the nest materials. Three nests hosted two species of tineid moths: Tinea translucens (Fig. 2e) and Niditinea baryspilas DISCUSSION (Table 3). The nest detritus consisted of decaying wood chips, a lot of down, pellets, food remains Owls feed on a wide range of prey including small (crushed insects) and droppings. mammals, birds, fish and insects, depending on the owl species, and carry similar items back to their 4) Collared Scops Owl and Ryukyu Scops Owl nests as food for their nestlings. As a result protein, Twelve nests supported four tineid species: Opog- in the form of keratin (feathers, down, fur, crushed ona sacchari (Fig. 2h), O. sp. (Fig. 2j), Phaeoses sp. feather sheaths, pellets, food remains) or chitin (Fig. 2k) and Setomorpha sp. (Fig. 2l), of which O. (arthropod exoskeleton material) is typically present sacchari predominated (Table 4). From one nest, one in owl nests, and is considered to be an abundant

81 Y. NASU et al.

Table 5. Relationships among nest detritus and food habits of Lepidoptera in owl nests in Japan.

Keratinophagous, chitinophagous, coprophagous Detritophagous Phytophagous, polyphagous

Dead plant Keratin,* droppings Chitin,** keratin,* droppings Dead plant matter, dried plants Living plants Dead leaves? Bee nest Unkown Animal matter matter derived from bird Owls Locality Food of owl breeding in the Tineidae Oecophoridae Elachistidae Crambidae Pyralidae nest Monopis Monopis Monopis Monopis Tinea Niditinea Niditinea Niditinea Opogona Opogona Phaeoses Setomorpha Martyringa Agonopterix Mabra Pyralis Endotricha longella flavidorsalis sp. congestella translucens baryspilas striolella sp. sacchari sp. sp. sp. ussuriella sp. charonialis regalis theonalis (= pavlovskii)

Blakiston’s Fish Owl, Keratin,* Hokkaido Fish ○ ○ Ketupa blakistoni droppings Ural Owl, Strix Mouse, small bird, Keratin,* Honshu, Kyushu ◎ ◎ ○ ◎ ◎ ◎ ○ ○ ○ ○ uralensis small animal droppings Brown Hawk-Owl, Chitin,** keratin,* Honshu, Kyushu Insect ○ ◎ Ninox scutulata droppings Collared Scops Owl, Insect, small Chitin,** keratin,* Ryukyu (Okinawa Is.) ◎ ○ ○ Otus lempiji animal droppings Ryukyu Scops Owl, Ryukyu (Okinawa Is., Chitin,** keratin,* Insect ◎ ○ ○ ○ Otus elegans Minami-daito Is.) droppings

* down, fur, crushed feather sheaths, pellets; ** pellets, crushed insects; *** adults hibernating. ◎: dominant. source of food especially for tineids, with some spe- was reared had many dead moss and liverwort, which cies being keratinophagous and others chitinopha- were considered to be gathered by tits after the nest gous (Robinson & Nielsen 1993; Nasu et al. 2007a). had been abandoned halfway through breeding, and the owl nest from which one crambid moth was 1) Lepidoptera fauna of owl nests reared had a lot of bark torn into ribbons, which were During the course of this study, Lepidoptera from considered to be gathered by Japanese Giant Flying Blakiston’s Fish Owl, Brown Hawk-Owl, Collared Squirrel, Petaurista lecogenys. Pyralid moth larvae Scops Owl and Ryukyu Scops Owl nests were newly typically feed in bee nests, while crambid moth lar- recorded, and five species (Niditinea striolella, vae feed on dead leaves (Inoue 1982). In these two Agonopterix sp., Mabra charonialis, Pyralis regalis nests, the food of the pyralid moth is unclear, how- and Endotricha theonalis) were recorded from bird ever the crambid moth larva is presumed to have fed nests for the first time (see Table 5). on plant matter. N. striolella developed from a Blaksiton’s Fish Tinea translucens and Niditinea baryspilas both Owl nest in February, revealing that either the larvae developed from Brown Hawk-Owl nests. The former or the pupae had hibernated in the nest. The moth species is known as a household pest and feeds on larva is suspected of having fed on owl down, because fur. In nature, the moth has been found in bird nests related Niditinea species are chitinophagous (Eurasian Tree Sparrow, Passer montanus, Red- (Robinson & Nielsen 1993). Adult Agonopterix sp. rumped Swallow, Hirundo daurica, and Barn Swal- was found in the nest in October, and the larvae had low, H. rustica) breeding in urban areas (Kiritani not developed and the adults possibly hibernated in 1959; Tomioka & Nakamura 2000; Nasu et al. the nest because all known congeners are herbivo- 2007b). T. translecens is considered to feed on down rous. The small number of moths found in Blakiston’s in nests, and N. baryspilas on chitin. T. translecens Fish Owl nests was possibly related to the fact that was found only in Brown Hawk-Owl nests restricted there was little natural detritus and few prey remains to urban areas. in the nests. Several tineid moths developed from Collared Many tineid species developed from Ural Owl Scops Owl and Ryukyu Scops Owl nests in Okinawa nests, with keratinophagous Monopis spp. and chitin- Prefecture. The nests contained considerable amounts ophagous Nidititea spp. predominating. The tineid of chitin (Collared Scops Owl feeds on insects and species found did not vary regionally. Many M. lon- small animals, and Ryukyu Scops Owl on insects). gella moths were reared from larvae hibernating in Opogona sacchari is known to be a pest of many nest detritus. The nest from which one pyralid moth tropical crops, fruits and ornamental plants

82 Nest-dwelling Lepidoptera and owls

Table 5. Relationships among nest detritus and food habits of Lepidoptera in owl nests in Japan.

Keratinophagous, chitinophagous, coprophagous Detritophagous Phytophagous, polyphagous

* down, fur, crushed feather sheaths, pellets; ** pellets, crushed insects; *** adults hibernating. ◎: dominant.

(Yoshimatsu et al. 2004); however, in nests it is even wider range of animals live in nests and form a thought to feed on decaying wood chips in the detri- complex food web there. tus of natural nests and in nest boxes. The food of Phaeoses is unknown, but among related genera the 2) Role of owls as ecosystem engineers and Lep- Opogona are scavengers, and the Setomorpha feed idoptera on dried plants (Robinson & Nielsen 1993). Endotri- Owls utilize natural hollow trees or nest boxes for cha theonalis may feed on decaying plants in nests, breeding, and gather no nest materials (Nakamura & because it is known to feed on old leaves (Inoue Nakamura 1995; Yuko Hayashi personal communica- 1982). tion); however, debris (down, feathers, fur, crushed Keratinophagous M. longella, which predominated feather sheaths, pellets) and droppings derived from in Ural Owl nests in Honshu, has not been found in juvenile and parent owls accumulate at the bottom of scops owl nests, although the moth is also found in nest cavities as a bed on which eggs and chicks rest, Okinawa Prefecture. This is likely to be because the and incidentally provide important food for tineid scops owl species of Okinawa are largely insectivo- moths. Such keratin sources are typically heavily rous hence their nests contain mostly chitin, not ker- soiled by juvenile owl droppings, and tineid larvae atin; however, chitinophagous Niditinea spp., widely seem to prefer soiled keratin rather than those free of distributed in Asia, also were not reared from the organic contamination. When fed on uncontaminated Okinawan scops owl nests, and the reasons for this feathers or animal hair, their growth rate is much are unknown. slower and mortality much higher than on other diets Many tineid moth species were found to utilize (Hinton 1956; Robinson 1979, 1988, 1990). Owls owl nests, with the moth fauna varying among the tend to re-utilize the same nests for breeding in con- owl species. The differences appear to be related to secutive years (Nakamura & Nakamura 1995; Yuko the food habits of the owl hosts and the owl habitats, Hayashi, Aki Higuchi, Masaoki Takagi personal with tineid species specifically selecting certain types communications), thus there is multi-year accumula- of owl nests. Beetles, Trox nohirai and Protaetia tion of debris within their nesting cavities. These brevitarsis, are also known to live in Ural Owl nests accumulated keratin sources provide considerable where they serve as important decomposers of nest suitable food for tineid moths in owl nests, thus the detritus (Inagaki 2009; Nasu et al. 2011). Further- owls play an important role in providing suitable more, many species of ants and flies are found in owl habitat and optimal food for tineids. As Nasu et al. nests (Yoshitsugu Nasu and Syo Daimon, unpub- (2007a) have already stated, tineids are thought to lished data), and further studies may reveal that an help maintain the cleanliness of owl nest chambers

83 Y. NASU et al. by decomposing keratin sources. Such relationships Linn Soc 163: 1–14. between tineids and owls may be mutualistic. Further Inagaki M (2009) Trox nohirai Nakane. In: Tsukamoto quantitative studies on rates of decomposition of owl K, Inagaki M, Kawahara M & Mori M, Japanese nest detritus by tineids and of the interaction between coprophagous scarab-beetles. pp 58–60. Tonbo owls and tineids are necessary to further understand Press, Osaka (in Japanese). the extent of this mutualism. Inoue H (1982) Pyralidae. In: Inoue H, Sugi S, Kuroko Murahama et al. (2007) reported that the night- H, Moriuti S, Kawabe A & Owada M, Moths of time temperatures within Ural Owl nest boxes during Japan 1. pp 307–404, 2 pp 223–254, pls. 36–48, the incubation period in winter were always higher (a 228, 296, 298–314. Kodansha, Tokyo (in Japanese). maximum of 4.4°C) than the ambient temperature Jalava J (1980) Tineidae (Lepidoptera) from nests of the Ural Owl (Strix uralensis Pall.). Notulae Entomol 60: outside. Such nest warmth may encourage the devel- 96–100. opment of tineid larvae during this period. Jones CG, Lawton JH & Shachak M (1994) Organisms as ecosystem engineers. OIKOS 69: 373–386. 3) Adaptation of Lepidoptera to nest-dwelling Kiritani K (1959) Natural habitats of household insect Tineid moths reared from owl nest detritus fly pests . Shin-Konchu 12(2): 2–6 (in Japanese). poorly, fall on disturbance, crash-dive and burrow Murahama S, Nasu Y & Matsumuro H (2007) Monitor- into nest materials. Such behaviors are typically ing the breeding stage of the Ural owl by thermo found among tineids living in bird nests (Robinson recorder. Bird Research 3: T13–T19 (in Japanese 2004; Nasu et al. 2012). That tineids burrow rapidly with English summary). into nest materials may indicate that they have Nakamura T & Nakamura M (1995) Birds’ life in Japan evolved an escape strategy with which to avoid being with color pictures: birds of mountain, woodland and preyed upon by their nest hosts. field. Hoikusha, Osaka (in Japanese). Nasu Y, Huang GH, Murahama S & Hirowatari T (2008) Tineid moths (Lepidoptera, Tineidae) from Goshawk ACKOWLEDGMENTS and Ural Owl nests in Japan, with notes on larviparity Our thanks are due to the following naturalists for their of Monopis congestella (Walker). Trans lepid Soc kind advice and help in collecting owl nests: Kana Akatani, Japan 59: 187–193. Nobuyuki Azuma, Sho Daimon, Kenzo Fukuoka, Keitaro Nasu Y, Murahama S, Matsumuro H, Hashiguchi D & Harusawa, Daisuke Hashiguchi, Yuko Hayashi, Aki Higuchi, Murahama C (2007a) First record of Lepidoptera Ikuko Inamori, Teruaki Ishii, Masayuki Iwasaki, Utsugi Jinbo, from Ural owl nests in Japan. Appl Entomol Zool 42: Naoya Kumashiro, Chieko Murahama, Teiji Nakao, Tokio 607–612. Sugiyama, Ken Takenaka, Tosihisa Saito, Makoto Sakai, Manabu Shibao, Michiyo Takada, Masaoki Takagi, Kazuhiko Nasu Y, Murahama S, Matsumuro H, Ueda K & Takizawa, Hiroshi Tanaka,. Tomokazu Tanigawa, Takaaki Hirowatari T (2012) Lepidoptera from nests of four Tomosada, and Masahiro Toyama. This work was supported insectivorous bird species. Trans lepid Soc Japan 63: partially by Grants-in-Aid for Scientific Research (nos. 87–93 (in Japanese with English summary). 19916019, 21570102) from the Japan Ministry of Education, Nasu Y, Murahama S, Matsumuro H, Ueda K, Hirowatari Culture, Sports, Science and Technology. T & Yoshiyasu Y (2011) Protaetia brevitarsis (Lewis) (Coleoptera, Scarabaeidae) from the Ural Owl nests REFERENCES in Japan. Strix 27: 67–72 (in Japanese with English summary). Hicks EA (1959) Check-List and Bibliography on the Nasu Y, Murahama S, Sakai M & Yamauchi T (2007b) Occurrence of Insects in Birds’ Nests. The Iowa State Tineidae (Lepidoptera) from bird’s nests and pellets, College Press, Ames. and carnivore feces in Japan. Jpn J Entomol (New Hicks EA (1962) Check-list and bibliography on the Series) 10: 89–97 (in Japanese with English sum- occurrence of insects in birds’ nests. Suppl. 1. Iowa mary). State J Sci 36: 233–348. Nordberg S (1936) Biologisch-ökologische untersuc- Hinton HE (1956) The larvae of the species of Tineidae hungen über die Vogelnidicolen [Biological and eco- of economic importance. Bull Entomol Res 47: 251– logical examinations about bird nests and nidicolous 346. fauna]. Acta Zool Fenn 21: 1–168. Huang GH, Chen LS, Hirowatari T, Nasu Y & Wang M Robinson GS (1979) Clothes-moths of the Tinea pel- (2011) A revision of the Monopis monachella species lonella complex: a revision of the world’s species complex (Lepidoptera: Tineidae) from China. Zool J (Lepidoptera: Tineidae). Bull Brit Mus Nat Hist

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