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Research Article
Fauna of nocturnal moth species collected in a semi-natural grassland at Kanpu-zan in northern Japan
expand article infoMasaru Kamikura‡,§, Yuzu Sakata§
‡ Sugadaira Research Station, Mountain Science Center, University of Tsukuba, Nagano, Japan
§ Department of Biological Environment, Faculty of Bioresource Sciences, Akita Prefectural University, Akita, Japan
Open Access

Abstract

Semi-natural grasslands, which house species-rich ecosystems, have rapidly declined since the twentieth century due to land-use practices, such as agricultural intensification and abandonment. Owing to their diversity and known habitat associations, nocturnal moths are considered as one of the most suitable organisms to be studied for assessing the dynamics of species composition as a result of changes in landscape management of semi-natural grasslands. The present study provides the foremost description of nocturnal moth fauna of the semi-natural grassland at Kanpu-zan, northern Japan. Moth population data from 1987 were compared to the data collected in 2018 to evaluate the impact of decline in grasslands on species-richness. During the field sampling in 2018, a total of 226 nocturnal moth species were detected, which was nearly two-thirds of the number of species recorded in 1987, i.e. 396 species. The values obtained in 2018 were found to be nearly constant for different sites. For both periods, it was evident that moth fauna in Kanpu-zan mainly consisted of species that relied on woody plants. Amongst the species which were only recorded in 1987, 107 species were generalists that fed on plants that are commonly distributed in Kanpu-zan. No moth species were recorded that depended upon endangered or extinct plant food sources. Thus, it is unlikely that the decline in the number of moth species in Kanpu-zan was due to the loss in plant food sources. Our results suggest that environmental factors other than food plants may have caused decline and changes in nocturnal moth fauna. More studies on various organism fauna are needed for understanding the conservation of semi-natural grassland, considering that the loss of semi-natural grasslands is one of the major threats to biodiversity.

Keywords

arthropods, northern Japan, nocturnal moth, semi-natural grassland, species occurences

Introduction

Semi-natural grasslands house species-rich ecosystems including regional meta-communities by specialists of open habitats (Emanuelsson 2008, Pykälä et al. 2005, Alison et al. 2017). The persistence of semi-natural grassland depends on anthropogenic activities, such as mowing, grazing and burning (Suka et al. 2012, Dengler et al. 2014, Ushimaru et al. 2018). Recent studies have shown that the grasslands may have persisted during the Holocene in a natural state due to natural disturbances and severe environmental conditions (Nakahama et al. 2018, Ohwaki 2018). However, these semi-natural grasslands have shown a continuous and rapid decline since the twentieth century due to changes in land-use practices such as agricultural intensification and abandonment (Donald et al. 2001, Eriksson et al. 2002, Koyanagi and Furukawa 2013). As a result, many grassland species, including insects feeding on grassland plants and those inhabiting grassland habitat, are presently endangered due to the scarcity of natural grasslands in these regions (Maes and Van Dyck 2001, Habel et al. 2013). There is an urgent need to understand the changes in grassland fauna due to human activities.

Nocturnal moths are an ecologically important group of insects that play a key role in herbivory and pollination. They act as a food source for birds as well as potential indicators of ecosystem change across a wide variety of landscapes (Kitching et al. 2000, Summerville and Crist 2004, Macgregor et al. 2016). Declines in moth populations have been linked to habitat loss and agricultural intensification (Wenzel et al. 2006, Merckx et al. 2012). Owing to their diversity and known habitat associations, nocturnal moths are considered as one of the most suitable study taxa for assessing species richness against the changes in landscape management of semi-natural grasslands (Erhardt and Thomas 1991, Wenzel et al. 2006; Alison et al. 2017).

Kanpu-zan, which is located in northern Japan, consists of semi-natural grasslands partly maintained by mowing and burning and accommodates many endangered grassland species (Akita Prefecture 2016). The region was a rich source of grassland resources as a livestock feed for farmers until the 1960s. However, the utilisation of the grassland, such as the grass harvest, gradually declined leading to succession by dense shrubs and trees from the 1970s as lifestyles changed. A study indicates that the grassland area, which was 319 ha in 1975, had decreased to 138 ha by 2014 (Masui et al. 2017). The prevalent plant community varied depending on the management frequency and plant richness was lower in abandoned sites with reduced or diminished grassland species (Masui et al. 2017). Documentation of moth species in the region was conducted in 1987 before the grassland had severely declined (Takahashi 1993). However, the present status of moth fauna is largely unknown. This study presents the first account of nocturnal moths in Kanpu-zan after the decline in grasslands and presents a model for understanding the changes in moth fauna related to altered vegetation in grassland areas caused by changes in human activity. Uchida and Ushimaru (2014) compared plant richness and butterfly and orthopteran richness and diversity amongst three different land use types in semi-natural grasslands: abandoned, traditional and intensified terraces and suggested that the number of individuals of most herbivorous species decreased randomly with respect to life-history traits, following a decline in plant richness reflecting disturbance frequency. Due to the present plant flora being revealed in Kanpu-zan, this site provides a promising opportunity to understand the relationship between moth fauna and the land use of the grassland vegetation.

Materials and Methods

Study site

Surveys of nocturnal moths were carried out at Kanpu-zan during May to October in 2018, twice a month (12 nights in total), on clear nights from sunset to midnight. Three sites that differ in vegetation were selected: 1) grassland maintained by mowing once or twice a year (grassland site), 2) site where grassland management had been abandoned leading to overgrowth of shrubs and trees (forest site) and 3) mixed vegetation with both grassland and woody plants (mixed site) (Fig. 1). In 1987, the moths were collected during May to September, once or twice a month (6 nights in total) at one site in a grassland dominated by tall grass such as Miscanthus sinensis, where shrubs and hedgerow trees were interspersed (Takahashi 1993).

Figure 1.  

Studied area and habitat vegetation: a. Geographic location of Kanpu-zan; b. grassland (GL); c. forest (FR); d. mixed vegetation (MIX).

Sampling methods

Moths were collected using UV light traps, equipped with a 20 W black (ultraviolet) light fluorescent tube. One light trap was set at each site. The same method was used in 1987 (Takahashi 1993). All collected specimens were dried at room temperature and mounted for morphological examination. All moths were identified to species level by M. Kamikura and K. Umetsu using descriptions and photos in the book "Standard of Moths in Japan" (Hirowatari et al. 2013, Kishida 2011a, Kishida 2011b, Nasu et al. 2013). Some of the family names (i.e. Arctiidae, Lymantriidae, Micronoctuidae) were updated to current taxonomy. All the specimens were preserved as a personal collection of M. Kamikura.

Data analyses

Owing to the annual variations in abundance of moth species (Highland et al. 2013), data analyses were based on presence-absence data of moths sampled at each site in each month (Table 1,Suppl. material 1). Species composition was compared between years and amongst sites. In addition, because micro-moths belonging to the families of Chrysopeleiidae, Cosmopterigidae, Oecophoridae and Depressariidae were not identified in the study in 1987 (Takahashi 1993), those specimens were excluded from the data in 2018, as well as from the following analysis. In order to test whether the community composition of moth species differed between 1987 and 2018, permutational multivariate analyses of variance (PERMANOVA) were performed. We included the year of the survey as an explanatory variable. Additionally, PERMANOVA was also performed to test whether the community composition of moth species differed amongst the three sites in 2018. Here each site was included as an explanatory variable. For these analyses, the ADONIS function in library VEGAN (Oksanen et al. 2016) in R. ver. 3.3.2 (R Development Core Team 2016) was used. Non-metric multidimensional scaling analysis (NMDS) on the Jaccard index was used to visually summarise the plant community compositions of each site at each month.

Species list (presence (1) or absence (0)) of the three sites in 2018 and one site in 1987.

family

species name

food

specialist/generalist

1987

forest

grassland

mixed

Bombycidae

Bombyx mandarina (Moore, 1872)

woody

specialist

0

1

1

0

Chrysopeleiidae

Ascalenia sp. (Japanese name: Zugurokobukazariba)

unknown

unknown

0

0

0

1

Cosmopterigidae

Pyroderces sarcogypsa (Meyrick, 1932)

unknown

unknown

0

0

0

1

Cosmopterigidae

Ressia quercidentella Sinev, 1988

unknown

unknown

0

1

0

1

Cossidae

Pharagmataecia castaneae (Hübner, 1790)

herb

specialist

1

0

0

0

Cossidae

Zeuzera multistrigata Moore, 1881

woody

generalist

0

1

1

1

Crambidae

Agrotera posticalis Wileman, 1911

woody

specialist

0

0

0

1

Crambidae

Anania albeoverbascalis Yamanaka, 1966

unknown

unknown

1

0

0

0

Crambidae

Anania verbascalis ([Denis & Schiffermüller], 1775)

herb

specialist

1

0

0

0

Crambidae

Ancylolomia japonica Zeller, 1877

herb

specialist

0

0

0

1

Crambidae

Bradina angustalis Yamanaka, 1984

unknown

unknown

1

0

0

0

Crambidae

Bradina atopalis (Walker, 1859)

unknown

unknown

0

1

1

1

Crambidae

Calamotropha okanoi Bleszynski, 1961

unknown

unknown

1

0

0

0

Crambidae

Camptomastix hisbonalis (Walker, 1859)

unknown

unknown

0

1

0

0

Crambidae

Chilo luteellus (Motschulsky, 1866)

herb

generalist

0

0

0

1

Crambidae

Chilo suppressalis (Walker, 1863)

herb

specialist

1

0

0

0

Crambidae

Chrysoteuchia diplogramma (Zeller, 1863)

herb

specialist

0

1

0

0

Crambidae

Circobotys nycterina Butler, 1879

unknown

unknown

1

0

0

0

Crambidae

Cnaphalocrocis medinalis (Guenée, 1854)

herb

specialist

1

0

0

0

Crambidae

Cnaphalocrocis stereogona (Meyrick, 1886)

herb

specialist

1

1

0

0

Crambidae

Crambus argyrophorus Butler, 1878

unknown

unknown

1

0

0

0

Crambidae

Crambus perlellus (Scopoli, 1763)

herb

specialist

1

0

0

0

Crambidae

Daulia afralis Walker, 1859

herb

specialist

1

0

0

0

Crambidae

Diasemia reticularis (Linnaeus, 1761)

herb

generalist

1

0

0

0

Crambidae

Dolicharthria bruguieralis (Duponchel, 1833)

herb

generalist

1

0

0

0

Crambidae

Flavocrambus striatellus (Leech, 1889)

unknown

unknown

1

0

0

0

Crambidae

Glaucocharis exsectella (Christoph, 1881)

lichen/moss

specialist

1

0

0

1

Crambidae

Glyphodes pryeri Butler, 1879

woody

specialist

1

0

0

0

Crambidae

Haritalodes derogatus (Fabricius, 1775)

woody

specialist

0

0

0

1

Crambidae

Herpetogramma luctuosale (Guenée, 1854)

woody

specialist

1

1

1

1

Crambidae

Herpetogramma magnum (Butler, 1879)

herb/woody

generalist

0

0

0

1

Crambidae

Herpetogramma moderatale (Christoph, 1881)

herb

generalist

1

0

0

0

Crambidae

Herpetogramma stultale (Walker, 1859)

herb

generalist

1

0

0

0

Crambidae

Mabra charonialis (Walker, 1859)

dead leaves

generalist

1

0

0

0

Crambidae

Maruca vitrata (Fabricius, 1787)

herb

specialist

1

0

0

0

Crambidae

Microchilo inouei Okano, 1962

unknown

unknown

1

0

0

0

Crambidae

Nacoleia commixta (Butler, 1879)

woody

generalist

1

1

0

0

Crambidae

Nacoleia sibirialis (Millière, 1879)

unknown

unknown

1

0

0

0

Crambidae

Nacoleia tampiusalis (Walker, 1859)

unknown

unknown

0

0

1

1

Crambidae

Nomis albopedalis Motschulsky, 1861

unknown

unknown

1

1

1

1

Crambidae

Nomophila noctuella (Denis & Schiffermüller, 1775)

herb/woody

generalist

1

0

0

0

Crambidae

Omiodes tristrialis (Bremer, 1864)

unknown

unknown

1

0

0

1

Crambidae

Ostrinia latipennis (Warren, 1892)

herb

specialist

1

0

0

0

Crambidae

Palpita nigropunctalis (Bremer, 1864)

woody

generalist

0

1

1

1

Crambidae

Patissa fulvosparsa (Butler, 1881)

unknown

unknown

0

0

0

1

Crambidae

Piletocera sodalis (Leech, 1889)

unknown

unknown

1

0

0

0

Crambidae

Platytes ornatella (Leech, 1889)

unknown

unknown

1

0

0

0

Crambidae

Pleuroptya chlorophanta (Butler, 1878)

woody

generalist

1

0

0

0

Crambidae

Pleuroptya harutai (Inoue, 1955)

woody

specialist

0

0

1

0

Crambidae

Pleuroptya inferior (Hampson, 1898)

herb

specialist

1

0

0

0

Crambidae

Prodasycnemis inornata (Butler, 1879)

herb

specialist

1

0

0

0

Crambidae

Pyrausta unipunctata Butler, 1881

unknown

unknown

1

0

0

0

Crambidae

Scirpophaga virginia Schultze, 1908

unknown

unknown

1

0

0

0

Crambidae

Sitochroa palealis (Denis & Schiffermüller, 1775)

herb

specialist

0

0

1

0

Crambidae

Sitochroa umbrosalis (Warren, 1892)

herb

specialist

1

1

1

1

Crambidae

Sitochroa verticalis (Linnaeus, 1758)

herb

generalist

0

0

1

1

Crambidae

Spoladea recurvalis (Fabricius, 1775)

herb

generalist

1

1

1

0

Crambidae

Syllepte pallidinotalis (Hampson, 1912)

woody

generalist

1

0

0

0

Crambidae

Syllepte segnalis (Leech, 1889)

unknown

unknown

1

0

0

0

Crambidae

Torulisquama obliquilinealis Inoue, 1982

unknown

unknown

1

0

0

0

Crambidae

Tyspanodes striatus (Butler, 1879)

woody

generalist

1

0

0

0

Crambidae

Udea lugubralis (Leech, 1889)

unknown

unknown

1

0

0

0

Crambidae

Uresiphita gracilis (Butler, 1879)

unknown

unknown

1

0

0

0

Crambidae

Xanthocrambus lucellus (Herrich-Schäffer, 1848)

unknown

unknown

0

1

1

0

Depressariidae

Agonopterix mutuurai Saito, 1980

herb

specialist

0

0

0

1

Depressariidae

Agonopterix sumizome Fujisawa, 1985

herb

specialist

0

0

0

1

Depressariidae

Agonopterix yomogiella Saito, 1980

herb

specialist

0

0

0

1

Drepanidae

Habrosyne pyritoides (Butler, 1766)

woody

specialist

1

0

0

0

Drepanidae

Tethea ampliata (Butler, 1878)

woody

specialist

1

0

0

0

Drepanidae

Thyatira batis (Linnaeus, 1758)

woody

specialist

1

0

0

0

Erebidae

Arctornis l-nigrum (Müller, 1764)

woody

generalist

0

1

0

0

Erebidae

Artaxa subflava (Bremer, 1864)

herb/woody

generalist

1

1

1

1

Erebidae

Barsine aberrans (Butler, 1877)

lichen/moss

generalist

0

1

0

0

Erebidae

Barsine pulchra (Butler, 1877)

unknown

unknown

1

0

1

0

Erebidae

Barsine striata (Bremer & Grey, 1853)

dead leaves

generalist

1

0

0

1

Erebidae

Chinoarctia nivea (Ménétriés, 1859)

herb

generalist

1

1

1

1

Erebidae

Cifuna locuples Walker, 1855

herb/woody

generalist

1

0

0

0

Erebidae

Cyana hamata (Walker, 1854)

lichen/moss

generalist

1

0

0

1

Erebidae

Eilema affineola (Bremer, 1864)

unknown

unknown

1

1

1

1

Erebidae

Eilema deplena (Esper, 1787)

lichen/moss

generalist

1

1

0

0

Erebidae

Eilema japonica (Leech, 1889)

lichen/moss

generalist

0

1

0

0

Erebidae

Eilema nankingica (Daniel, 1954)

unknown

unknown

1

0

0

0

Erebidae

Eilema vetusta (Walker, 1854)

lichen/moss

generalist

1

1

0

1

Erebidae

Epatolmis caesarea (Goeze, 1781)

herb/woody

generalist

1

1

1

0

Erebidae

Ghoria collitoides Butler, 1885

unknown

unknown

1

1

1

1

Erebidae

Ghoria gigantea (Oberthür, 1879)

woody

generalist

0

1

0

1

Erebidae

Ilema eurydice (Butler, 1885)

woody

generalist

1

0

1

0

Erebidae

Kidokuga piperita (Oberthür, 1880)

woody

generalist

1

0

0

1

Erebidae

Lemyra imparilis (Butler, 1877)

woody

generalist

1

0

0

0

Erebidae

Lemyra inaequalis (Butler, 1879)

woody

generalist

1

0

0

0

Erebidae

Lithosia quadra (Linnaeus, 1758)

lichen/moss

generalist

1

0

0

0

Erebidae

Lymantria mathura (Moore, 1865)

woody

generalist

1

1

0

0

Erebidae

Miltochrista calamina Butler, 1877

woody

generalist

1

1

1

1

Erebidae

Miltochrista miniata (Forster, 1771)

lichen/moss

generalist

1

1

0

1

Erebidae

Mimachrostia fasciata Sugi, 1982

unknown

unknown

1

1

0

0

Erebidae

Orgyia thyellina Butler, 1881

herb/woody

generalist

0

1

0

0

Erebidae

Pelosia muscerda (Hufnagel, 1766)

lichen/moss

generalist

1

0

0

1

Erebidae

Pelosia noctis (Butler, 1881)

lichen/moss

generalist

1

0

0

0

Erebidae

Pelosia obtusa (Herrich-Schäffer, 1852)

unknown

unknown

1

0

0

0

Erebidae

Phragmatobia amurensis Seitz, 1910

herb

generalist

1

0

0

0

Erebidae

Rhyparioides amurensis (Bremer, 1861)

herb

specialist

1

1

1

1

Erebidae

Rhyparioides metelkana (Lederer, 1861)

herb

generalist

1

0

1

0

Erebidae

Rhyparioides nebulosa Butler, 1877

herb

generalist

1

1

0

1

Erebidae

Sphrageidus similis (Fuessly, 1775)

woody

generalist

1

1

0

0

Erebidae

Spilarctia lutea (Hufnagel, 1766)

unknown

unknown

1

0

0

0

Erebidae

Spilarctia obliquizonata (Miyake, 1910)

woody

specialist

0

1

0

0

Erebidae

Spilarctia seriatopunctata (Motschulsky, 1861)

woody

generalist

1

0

1

1

Erebidae

Spilosoma lubricipedum (Linnaeus, 1758)

woody

generalist

1

0

0

0

Erebidae

Spilosoma punctarium (Stoll, 1782)

woody

generalist

1

1

1

0

Erebidae

Stigmatophora rhodophila (Walker, 1865)

lichen/moss

generalist

1

0

0

0

Erebidae

Thumatha ochracea (Bremer, 1861)

unknown

unknown

1

0

0

0

Eupterotidae

Apha aequalis (Felder, 1874)

herb/woody

generalist

1

1

1

1

Geometridae

Abraxas fulvobasalis Warren, 1894

woody

specialist

1

0

0

0

Geometridae

Abraxas miranda Butler, 1878

woody

specialist

1

0

0

0

Geometridae

Abraxas niphonibia Wehrli, 1935

woody

specialist

1

0

1

0

Geometridae

Acrodontis fumosa (Prout, 1930)

woody

specialist

0

0

0

1

Geometridae

Amraica superans (Butler, 1878)

woody

specialist

1

1

1

0

Geometridae

Arichanna melanaria (Linnaeus, 1758)

woody

specialist

1

0

0

0

Geometridae

Ascotis selenaria (Denis & Schiffermüller, 1775)

herb/woody

generalist

1

0

0

0

Geometridae

Astygisa chlororphnodes (Wehrli, 1936)

woody

specialist

1

1

0

0

Geometridae

Chariaspilates formosaria (Eversmann, 1837)

herb/woody

generalist

0

1

1

1

Geometridae

Chiasmia defixaria (Walker, 1861)

woody

specialist

0

0

1

0

Geometridae

Chiasmia hebesata (Walker, 1861)

herb/woody

generalist

1

0

1

1

Geometridae

Chlorissa amphitritaria (Oberthür, 1879)

woody

generalist

1

0

0

1

Geometridae

Chlorissa anadema (Prout, 1930)

herb/woody

generalist

1

0

0

0

Geometridae

Chlorissa obliterata (Walker, 1863)

herb

generalist

1

0

0

0

Geometridae

Chloroclystis v-ata (Haworth, 1809)

herb/woody

generalist

1

1

0

0

Geometridae

Cleora insolita (Butler, 1878)

woody

generalist

1

0

0

0

Geometridae

Comibaena delicatior (Warren, 1897)

woody

generalist

1

0

0

1

Geometridae

Costaconvexa caespitaria (Christoph, 1881)

unknown

unknown

0

0

0

1

Geometridae

Culpinia diffusa (Walker, 1861)

herb/woody

generalist

1

0

0

0

Geometridae

Cusiala stipitaria (Oberthür, 1880)

woody

generalist

1

0

0

0

Geometridae

Descoreba simplex Butler, 1878

woody

generalist

1

0

0

0

Geometridae

Dindica virescens (Butler, 1878)

woody

specialist

1

1

0

0

Geometridae

Ecliptopera capitata (Herrich-Schäffer, 1840)

woody

specialist

1

0

0

0

Geometridae

Ecliptopera umbrosaria (Motschulsky, 1861)

woody

specialist

1

0

0

0

Geometridae

Ecpetelia albifrontaria (Leech, 1891)

woody

specialist

1

0

0

0

Geometridae

Ectropis excellens (Butler, 1884)

herb/woody

generalist

1

0

0

0

Geometridae

Ectropis obliqua (Prout, 1915)

herb/woody

generalist

1

0

0

0

Geometridae

Epirrhoe supergressa (Butler, 1878)

herb

specialist

1

0

0

0

Geometridae

Euphyia cineraria (Butler, 1878)

unknown

unknown

1

0

0

0

Geometridae

Eupithecia interpunctaria Inoue, 1979

herb

specialist

1

0

0

0

Geometridae

Eupithecia mandschurica Staudinger, 1897

unknown

unknown

1

0

0

0

Geometridae

Eupithecia neosatyrata Inoue, 1979

woody

specialist

1

0

0

0

Geometridae

Evecliptopera illitata (Wileman, 1911)

woody

specialist

1

0

1

1

Geometridae

Gandaritis fixseni (Bremer, 1864)

woody

generalist

0

0

0

1

Geometridae

Geometra dieckmanni Graeser, 1889

woody

specialist

0

0

0

1

Geometridae

Heterophleps fusca (Butler, 1878)

unknown

unknown

0

1

0

0

Geometridae

Heterothera postalbida (Wileman, 1911)

woody

specialist

1

0

0

0

Geometridae

Hypomecis punctinalis (Scopoli, 1763)

woody

generalist

1

0

0

0

Geometridae

Idaea auricruda (Butler, 1879)

unknown

unknown

1

1

0

1

Geometridae

Idaea denudaria (Prout, 1913)

unknown

unknown

1

0

0

0

Geometridae

Idaea invalida (Butler, 1879)

unknown

unknown

1

0

0

0

Geometridae

Idaea jakima (Butler, 1878)

unknown

unknown

1

0

0

0

Geometridae

Idaea muricata (Hufnagel, 1767)

unknown

unknown

1

1

0

1

Geometridae

Idaea nielseni (Hedemann, 1879)

unknown

unknown

0

0

1

0

Geometridae

Idaea trisetata (Prout, 1922)

unknown

unknown

1

0

0

0

Geometridae

Jankowskia pseudathleta Sato, 1980

woody

generalist

1

0

0

0

Geometridae

Lamprocabera candidaria (Leech, 1897)

woody

specialist

0

1

0

1

Geometridae

Lassaba nikkonis (Butler, 1881)

woody

generalist

1

0

0

0

Geometridae

Lomographa bimaculata (Fabricius, 1775)

woody

specialist

0

0

1

0

Geometridae

Lomographa temerata (Denis & Schiffermüller, 1775)

woody

specialist

1

1

1

1

Geometridae

Macaria liturata (Clerck, 1759)

woody

specialist

0

0

0

1

Geometridae

Macaria shanghaisaria Walker, 1861

woody

specialist

1

0

0

0

Geometridae

Menophra senilis (Butler, 1878)

herb/woody

generalist

1

1

0

0

Geometridae

Metabraxas clerica Butler, 1881

unknown

unknown

1

1

0

0

Geometridae

Microcalicha sordida (Butler, 1878)

woody

specialist

1

0

0

0

Geometridae

Ninodes splendens (Butler, 1878)

woody

specialist

1

0

0

0

Geometridae

Orthonama obstipata (Fabricius, 1794)

herb/woody

generalist

1

0

0

0

Geometridae

Otoplecta frigida (Butler, 1878)

woody

specialist

1

0

0

0

Geometridae

Ourapteryx maculicaudaria (Motschulsky, 1866)

woody

generalist

1

0

0

0

Geometridae

Oxymacaria normata (Alphéraky, 1892)

woody

specialist

1

0

0

0

Geometridae

Pachista superans (Butler, 1878)

woody

specialist

0

1

1

1

Geometridae

Pasiphila excisa (Butler, 1878)

woody

generalist

0

1

0

1

Geometridae

Peratophyga hyalinata (Kollar, 1844)

herb

specialist

1

0

0

0

Geometridae

Petelia rivulosa (Butler, 1881)

woody

specialist

1

0

0

0

Geometridae

Petrophora chlorosata (Scopoli, 1763)

herb

specialist

1

1

1

1

Geometridae

Phthonosema tendinosaria (Bremer, 1864)

herb/woody

generalist

1

1

1

1

Geometridae

Pingasa alba Swinhoe, 1891

woody

specialist

0

1

0

0

Geometridae

Pingasa pseudoterpnaria (Guenée, 1857)

woody

specialist

1

0

0

0

Geometridae

Protoboarmia faustinata (Warren, 1897)

woody

generalist

1

0

0

0

Geometridae

Protoboarmia simpliciaria (Leech, 1897)

woody

generalist

0

1

0

0

Geometridae

Pseuderannis lomozemia (Prout, 1930)

woody

generalist

1

0

0

0

Geometridae

Pylargosceles steganioides (Butler, 1878)

herb/woody

generalist

1

0

0

0

Geometridae

Racotis petrosa (Butler, 1879)

woody

specialist

1

0

0

0

Geometridae

Scopula floslactata (Haworth, 1809)

herb

generalist

0

1

0

0

Geometridae

Scopula ignobilis (Warren, 1901)

herb/woody

generalist

1

0

0

0

Geometridae

Scopula impersonata (Walker, 1861)

herb

specialist

1

0

0

0

Geometridae

Scopula nigropunctata (Hufnagel, 1767)

herb/woody

generalist

1

0

0

0

Geometridae

Scopula nupta (Butler, 1878)

unknown

unknown

1

0

0

0

Geometridae

Scopula pudicaria (Motschulsky, 1861)

herb

specialist

0

0

0

1

Geometridae

Scopula semignobilis Inoue, 1942

herb

specialist

1

0

0

0

Geometridae

Spilopera debilis (Butler, 1878)

woody

specialist

1

1

1

1

Geometridae

Trichopteryx ussurica (Wehrli, 1927)

unknown

unknown

0

1

0

0

Geometridae

Tyloptera bella (Butler, 1878)

woody

specialist

1

1

0

0

Geometridae

Xanthorhoe biriviata (Borkhausen, 1794)

herb

specialist

1

1

0

0

Geometridae

Xerodes albonotaria (Bremer, 1864)

herb/woody

generalist

1

0

0

0

Geometridae

Xerodes rufescentara (Motschulsky, 1861)

woody

generalist

1

0

0

0

Geometridae

Xerodes semilutata (Lederer, 1853)

woody

specialist

1

1

1

1

Lasiocampidae

Dendrolimus spectabilis (Butler, 1877)

woody

specialist

1

0

0

0

Lasiocampidae

Euthrix albomaculata (Bremer, 1861)

herb

specialist

1

0

0

0

Lasiocampidae

Euthrix potatoria (Linnaeus, 1758)

herb/woody

generalist

1

1

1

0

Lasiocampidae

Kunugia undans (Walker, 1859)

woody

generalist

0

1

1

1

Lasiocampidae

Malacosoma neustrium (Linnaeus, 1758)

woody

generalist

1

0

0

0

Limacodidae

Narosoideus flavidorsalis (Staudinger, 1887)

woody

generalist

1

1

0

1

Limacodidae

Parasa consocia Walker, 1863

woody

generalist

1

0

0

0

Limacodidae

Parasa hilarula (Staudinger, 1887)

woody

generalist

1

1

0

1

Limacodidae

Phlossa conjuncta (Walker, 1855)

unknown

unknown

1

0

0

0

Noctuidae

Abrostola ussuriensis Dufay, 1958

unknown

unknown

1

0

0

0

Noctuidae

Acanthoplusia agnata (Staudinger, 1892)

herb

generalist

0

0

1

0

Noctuidae

Acronicta hercules (Felder & Rogenhofer, 1874)

woody

specialist

0

0

1

0

Noctuidae

Acronicta rumicis (Linnaeus, 1758)

herb/woody

generalist

1

1

0

0

Noctuidae

Actinotia intermediata (Bremer, 1861)

herb

specialist

1

0

0

0

Noctuidae

Adrapsa simplex (Butler, 1879)

dead leaves

generalist

1

0

0

0

Noctuidae

Agrotis exclamationis (Linnaeus, 1758)

herb

generalist

1

0

1

1

Noctuidae

Agrotis ipsilon (Hufnagel, 1766)

herb/woody

generalist

0

0

1

0

Noctuidae

Agrotis segetum (Denis & Schiffermüller, 1775)

herb

generalist

1

0

0

0

Noctuidae

Amphipoea ussuriensis (Petersen, 1914)

unknown

unknown

1

0

0

0

Noctuidae

Amphipyra pyramidea (Linnaeus, 1758)

woody

generalist

1

0

0

0

Noctuidae

Anachrostis nigripunctalis (Wileman, 1911)

woody

specialist

1

1

0

0

Noctuidae

Anacronicta caliginea (Butler, 1881)

herb

specialist

1

0

0

0

Noctuidae

Anacronicta nitida (Butler, 1878)

herb

specialist

1

0

0

0

Noctuidae

Anapamea incerta (Staudinger, 1892)

unknown

unknown

1

0

0

0

Noctuidae

Anaplectoides virens (Butler, 1878)

herb

specialist

1

0

0

0

Noctuidae

Anarta (Calocestra) trifolii (Hufnagel, 1766)

herb

generalist

0

0

0

1

Noctuidae

Anorthoa angustipennis (Matsumura, 1926)

woody

generalist

1

0

0

0

Noctuidae

Apamea aquila (Donzel, 1837)

unknown

unknown

1

0

0

0

Noctuidae

Apamea hampsoni Sugi, 1963

unknown

unknown

1

1

0

0

Noctuidae

Apamea sordens (Hufnagel, 1766)

herb

specialist

0

0

0

1

Noctuidae

Archanara resoluta Hampson, 1910

herb

specialist

1

0

0

0

Noctuidae

Athetis albisignata (Oberthür, 1879)

herb

generalist

1

0

0

0

Noctuidae

Athetis cinerascens (Motschulsky, 1861)

unknown

unknown

1

0

0

0

Noctuidae

Athetis dissimilis (Hampson, 1909)

herb

generalist

1

0

0

0

Noctuidae

Athetis lapidea Wileman, 1911

unknown

unknown

1

1

1

1

Noctuidae

Athetis lineosa (Moore, 1881)

herb

generalist

1

0

0

0

Noctuidae

Athetis stellata (Moore, 1882)

herb

generalist

1

1

1

1

Noctuidae

Aventiola pusilla (Butler, 1879)

lichen/moss

generalist

1

0

0

0

Noctuidae

Axylia putris (Linnaeus, 1761)

herb

generalist

1

0

1

0

Noctuidae

Bambusiphila vulgaris (Butler, 1886)

herb

specialist

1

0

0

0

Noctuidae

Blasticorhinus ussuriensis (Bremer, 1861)

herb/woody

generalist

1

1

1

1

Noctuidae

Bomolocha melanica Sugi, 1959

unknown

unknown

1

0

0

0

Noctuidae

Bomolocha stygiana (Butler, 1878)

woody

specialist

1

0

0

1

Noctuidae

Bomolocha zilla (Butler, 1879)

unknown

unknown

1

0

0

0

Noctuidae

Callopistria juventina (Stoll, 1782)

herb

generalist

1

1

1

0

Noctuidae

Callopistria placodoides (Guenée, 1852)

herb

generalist

1

0

1

0

Noctuidae

Callopistria repleta Walker, 1858

herb

generalist

0

1

0

0

Noctuidae

Calyptra thalictri (Borkhausen, 1790)

herb

specialist

1

0

1

1

Noctuidae

Catocala duplicata Butler, 1885

woody

specialist

1

0

0

0

Noctuidae

Catocala nagioides Wileman, 1924

woody

specialist

1

0

0

0

Noctuidae

Catocala nubila Butler, 1881

woody

specialist

1

0

0

0

Noctuidae

Chandata bella (Butler, 1881)

herb

specialist

0

0

0

1

Noctuidae

Chasminodes sugii Kononenko, 1981

woody

specialist

1

0

0

0

Noctuidae

Chorsia japonica (Warren, 1912)

woody

specialist

0

0

0

1

Noctuidae

Chrysodeixis acuta (Walker, 1857)

herb/woody

generalist

1

0

0

0

Noctuidae

Chrysorithrum amatum (Bremer & Grey, 1853)

woody

specialist

1

0

0

1

Noctuidae

Chytonix subalbonotata Sugi, 1959

unknown

unknown

0

0

1

0

Noctuidae

Cidariplura gladiata Butler, 1879

unknown

unknown

1

0

0

0

Noctuidae

Clavipalpula aurariae (Oberthür, 1880)

woody

generalist

1

0

0

0

Noctuidae

Conistra grisescens Draudt, 1950

woody

generalist

1

0

0

0

Noctuidae

Corgatha nitens (Butler, 1879)

lichen/moss

generalist

0

1

0

0

Noctuidae

Cosmia trapezina (Linnaeus, 1758)

woody

generalist

1

0

0

0

Noctuidae

Cryphia mitsuhashi (Marumo, 1917)

lichen/moss

generalist

1

0

0

0

Noctuidae

Cucullia perforata Bremer, 1861

herb

specialist

0

1

0

0

Noctuidae

Diarsia canescens (Butler, 1878)

herb

generalist

1

1

1

1

Noctuidae

Diarsia deparca (Butler, 1879)

herb

generalist

0

0

0

1

Noctuidae

Diarsia pacifica Boursin, 1943

herb

generalist

1

0

0

0

Noctuidae

Diarsia ruficauda (Warren, 1909)

herb

generalist

1

0

0

0

Noctuidae

Diomea cremata (Butler, 1878)

lichen/moss

generalist

0

0

0

1

Noctuidae

Dipterygina cupreotincta Sugi, 1954

herb

specialist

1

0

0

0

Noctuidae

Dypterygia andreji Kardakoff, 1928

herb

specialist

1

0

0

0

Noctuidae

Ercheia niveostrigata Warren, 1913

herb

specialist

1

0

1

0

Noctuidae

Ercheia umbrosa Butler, 1881

woody

specialist

1

0

0

0

Noctuidae

Ericeia pertendens (Walker, 1858)

woody

specialist

0

1

0

0

Noctuidae

Erygia apicalis Guenée, 1852

herb/woody

specialist

0

0

1

0

Noctuidae

Euplexia lucipara (Linnaeus, 1758)

herb

generalist

0

1

1

0

Noctuidae

Eutelia geyeri (Felder & Rogenhofer, 1874)

woody

generalist

0

0

1

0

Noctuidae

Euxoa karschi (Graeser, 1889)

herb

specialist

1

0

0

0

Noctuidae

Euxoa sibirica (Boisduval, 1832)

herb

generalist

1

0

0

0

Noctuidae

Gonitis mesogona (Walker, 1858)

woody

generalist

1

0

0

0

Noctuidae

Gortyna basalipunctata Graeser, 1889

unknown

unknown

0

1

1

0

Noctuidae

Gortyna fortis (Butler, 1878)

herb

generalist

1

0

0

0

Noctuidae

Gynaephila maculifera Staudinger, 1892

lichen/moss

generalist

1

0

0

0

Noctuidae

Harita belinda (Butler, 1879)

woody

specialist

1

0

0

0

Noctuidae

Heliothis maritima Graslin, 1855

herb

generalist

0

0

1

0

Noctuidae

Herminia arenosa Butler, 1878

dead leaves

generalist

1

0

0

0

Noctuidae

Herminia grisealis (Denis & Schiffermüller, 1775)

dead leaves

generalist

1

0

0

0

Noctuidae

Herminia tarsicrinalis (Knoch, 1782)

dead leaves

generalist

1

0

0

0

Noctuidae

Holocryptis ussuriensis (Rebel, 1901)

unknown

unknown

1

0

0

0

Noctuidae

Honeyania ragusana (Freyer, 1844)

unknown

unknown

1

0

0

0

Noctuidae

Hydraecia petasitis Doubleday, 1847

herb

generalist

1

0

0

0

Noctuidae

Hydrillodes morosa (Butler, 1879)

dead leaves

generalist

1

1

1

1

Noctuidae

Hypena amica (Butler, 1878)

herb

specialist

0

1

0

0

Noctuidae

Hypena kengkalis Bremer, 1864

woody

specialist

1

0

0

0

Noctuidae

Hypena sp.

unknown

unknown

0

0

1

1

Noctuidae

Hypena tristalis Lederer, 1853

herb/woody

generalist

1

1

1

1

Noctuidae

Hypena whitelyi (Butler, 1879)

unknown

unknown

1

0

0

0

Noctuidae

Hyperstrotia flavipuncta (Leech, 1889)

dead leaves

generalist

1

1

1

1

Noctuidae

Iambia japonica Sugi, 1958

unknown

unknown

1

1

0

0

Noctuidae

Idia curvipalpis (Butler, 1879)

dead leaves

generalist

1

0

0

0

Noctuidae

Koyaga falsa (Butler, 1885)

herb

specialist

0

0

1

1

Noctuidae

Koyaga numisma (Staudinger, 1888)

unknown

unknown

1

0

0

0

Noctuidae

Leucapamea kawadai (Sugi, 1955)

unknown

unknown

1

0

0

0

Noctuidae

Lygephila maxima (Bremer, 1861)

herb

generalist

1

0

0

0

Noctuidae

Mamestra brassicae (Linnaeus, 1758)

herb

generalist

1

0

0

0

Noctuidae

Melapia electaria (Bremer, 1864)

herb

specialist

1

0

0

0

Noctuidae

Metopta rectifasciata (Ménétriès, 1863)

herb

specialist

1

1

0

1

Noctuidae

Micardia argentata Butler, 1878

herb

specialist

0

1

0

0

Noctuidae

Micreremites pyraloides Sugi, 1982

unknown

unknown

1

0

0

0

Noctuidae

Microxyla confusa (Wileman, 1911)

herb

specialist

1

0

0

0

Noctuidae

Mocis ancilla (Warren, 1913)

herb

specialist

1

0

0

0

Noctuidae

Mocis annetta (Butler, 1878)

herb/woody

specialist

1

0

1

0

Noctuidae

Mosopia sordidum (Butler, 1879)

unknown

unknown

1

0

0

0

Noctuidae

Mythimna chosenicola (Bryk, 1949)

unknown

unknown

1

0

0

0

Noctuidae

Mythimna flavostigma (Bremer, 1861)

unknown

unknown

1

0

1

1

Noctuidae

Mythimna inanis (Oberthür, 1880)

unknown

unknown

1

1

0

1

Noctuidae

Mythimna matsumuriana (Bryk, 1949)

herb

specialist

1

0

0

0

Noctuidae

Mythimna obsoleta (Hübner, 1803)

unknown

unknown

1

0

0

0

Noctuidae

Mythimna pallens (Linnaeus, 1758)

herb

generalist

1

0

0

0

Noctuidae

Mythimna postica (Hampson, 1905)

herb

specialist

1

0

0

0

Noctuidae

Mythimna pudorina (Denis & Schiffermüller, 1775)

herb

generalist

0

1

0

0

Noctuidae

Mythimna radiata (Bremer, 1861)

unknown

unknown

1

0

0

0

Noctuidae

Mythimna rufipennis Butler, 1878

unknown

unknown

1

0

1

1

Noctuidae

Mythimna separata (Walker, 1865)

herb

generalist

1

0

0

0

Noctuidae

Mythimna turca (Linnaeus, 1761)

herb

generalist

1

1

1

1

Noctuidae

Naranga aenescens Moore, 1881

herb

specialist

1

0

0

0

Noctuidae

Niphonyx segregata (Butler, 1878)

herb

specialist

1

0

0

0

Noctuidae

Nycteola asiatica (Krulikowski, 1904)

woody

generalist

1

0

0

0

Noctuidae

Ochropleura plecta (Linnaeus, 1761)

herb

generalist

1

0

0

0

Noctuidae

Orthogonia sera Felder & Felder, 1862

herb

specialist

1

0

0

0

Noctuidae

Orthosia ella (Butler, 1878)

herb

generalist

1

0

0

0

Noctuidae

Orthosia gothica (Linnaeus, 1758)

woody

generalist

0

1

0

0

Noctuidae

Orthosia lizetta Butler, 1878

woody

specialist

1

0

0

0

Noctuidae

Oruza mira (Butler, 1879)

dead leaves

generalist

1

0

0

0

Noctuidae

Pangrapta lunulata (Sterz, 1915)

woody

specialist

0

1

0

0

Noctuidae

Pangrapta obscurata (Butler, 1879)

woody

generalist

1

1

0

0

Noctuidae

Pangrapta umbrosa (Leech, 1900)

woody

specialist

1

0

1

0

Noctuidae

Panolis japonica Draudt, 1935

woody

specialist

1

0

0

0

Noctuidae

Paracolax trilinealis (Bremer, 1864)

unknown

unknown

1

0

0

0

Noctuidae

Paracolax tristalis (Fabricius, 1794)

dead leaves

generalist

0

1

0

1

Noctuidae

Paragabara flavomacula (Oberthür, 1880)

unknown

unknown

1

0

0

0

Noctuidae

Perigrapha hoenei Püngeler, 1914

woody

specialist

1

0

0

0

Noctuidae

Phyllophila obliterata (Rambur, 1833)

herb

specialist

1

1

1

1

Noctuidae

Plusiodonta casta (Butler, 1878)

woody

specialist

1

1

1

0

Noctuidae

Prospalta cyclica (Hampson, 1908)

unknown

unknown

1

0

0

0

Noctuidae

Protodeltote distinguenda (Staudinger, 1888)

herb

specialist

1

1

0

0

Noctuidae

Protodeltote pygarga (Hufnagel, 1766)

unknown

unknown

1

0

0

0

Noctuidae

Pyrrhia umbra (Hufnagel, 1766)

herb

generalist

1

0

0

0

Noctuidae

Rhizedra lutosa (Hübner, 1803)

herb

specialist

1

0

0

0

Noctuidae

Rivula sericealis (Scopoli, 1763)

herb

specialist

0

0

0

1

Noctuidae

Rusicada privata (Walker, 1865)

woody

specialist

0

0

1

0

Noctuidae

Sarcopolia illoba (Butler, 1878)

herb/woody

generalist

1

0

0

0

Noctuidae

Schrankia separatalis (Herz, 1905)

unknown

unknown

1

1

0

0

Noctuidae

Sesamia confusa (Sugi, 1982)

herb

specialist

1

0

0

1

Noctuidae

Sesamia turpis (Butler, 1879)

herb

specialist

1

0

0

0

Noctuidae

Sidemia bremeri (Erschoff, 1867)

unknown

unknown

0

1

1

0

Noctuidae

Sineugraphe bipartita (Graeser, 1889)

unknown

unknown

1

0

0

0

Noctuidae

Sineugraphe bipartita (Graeser, 1889)

unknown

unknown

0

1

0

0

Noctuidae

Sineugraphe exusta (Butler, 1878)

unknown

unknown

1

1

1

1

Noctuidae

Sineugraphe oceanica (Kardakoff, 1928)

herb

specialist

1

1

1

1

Noctuidae

Sophta subrosea (Butler, 1881)

unknown

unknown

1

0

0

0

Noctuidae

Sphragifera sigillata (Ménétriès, 1859)

woody

generalist

1

0

0

0

Noctuidae

Spodoptera depravata (Butler, 1879)

herb

specialist

1

0

0

0

Noctuidae

Stenhypena nigripuncta (Wileman, 1911)

lichen/moss

generalist

1

0

0

0

Noctuidae

Stenoloba clara (Leech, 1889)

unknown

unknown

0

0

1

0

Noctuidae

Stenoloba jankowskii (Oberthür, 1885)

unknown

unknown

0

0

0

1

Noctuidae

Sugia erastroides (Draudt, 1950)

unknown

unknown

1

0

0

0

Noctuidae

Sugia idiostygia (Sugi, 1958)

unknown

unknown

0

1

0

0

Noctuidae

Sugia stygia (Butler, 1878)

unknown

unknown

0

1

0

0

Noctuidae

Sypnoides picta (Butler, 1877)

woody

generalist

0

1

0

0

Noctuidae

Telorta edentata (Leech, 1889)

woody

generalist

0

0

1

0

Noctuidae

Thyas juno (Dalman, 1823)

woody

generalist

1

0

0

0

Noctuidae

Trachea punkikonis Matsumura, 1927

herb

specialist

1

0

0

0

Noctuidae

Trachea tokiensis (Butler, 1884)

herb

specialist

1

0

0

0

Noctuidae

Treitschkendia tarsipennalis (Treitschke, 1835)

dead leaves

generalist

1

0

0

0

Noctuidae

Virgo datanidia (Butler, 1885)

unknown

unknown

0

1

1

0

Noctuidae

Xestia c-nigrum (Linnaeus, 1758)

herb

generalist

1

0

0

0

Noctuidae

Xestia ditrapezium (Denis & Schiffermüller, 1775)

herb

generalist

1

0

0

0

Noctuidae

Xestia efforescens (Butler, 1879)

unknown

unknown

1

1

1

1

Noctuidae

Xestia stupenda (Butler, 1878)

herb

generalist

1

1

1

1

Noctuidae

Zanclognatha lunalis (Scopoli, 1763)

dead leaves

generalist

1

0

0

0

Noctuidae

Zekelita plusioides (Butler, 1879)

lichen/moss

generalist

1

0

0

0

Nolidae

Earias pudicana Staudinger, 1887

woody

generalist

1

1

0

0

Nolidae

Gelastocera exusta Butler, 1877

woody

generalist

1

0

0

0

Nolidae

Gelastocera kotschubeji Obraztsov, 1943

woody

generalist

1

0

0

0

Nolidae

Nola taeniata Snellen, 1874

unknown

unknown

0

1

0

0

Nolidae

Nola trilinea Marumo, 1923

herb

specialist

1

0

0

0

Nolidae

Pseudoips prasinanus (Linnaeus, 1758)

woody

specialist

1

0

1

0

Notodontidae

Clostera anachoreta Denis & Schiffermüller, 1775

woody

specialist

1

0

0

0

Notodontidae

Cutuza straminea (Walker, 1865)

herb

specialist

1

0

0

0

Notodontidae

Epodonta lineata (Oberthür, 1880)

woody

specialist

1

1

0

0

Notodontidae

Fentonia ocypete (Bremer, 1861)

woody

specialist

1

0

0

1

Notodontidae

Furcula furcula (Clerck, 1759)

woody

specialist

1

0

0

0

Notodontidae

Gonoclostera timoniorum (Bremer, 1861)

woody

specialist

1

0

0

0

Notodontidae

Hexafrenum leucodera (Staudinger, 1892)

woody

generalist

1

0

0

0

Notodontidae

Hupodonta corticalis Butler, 1877

woody

specialist

1

0

0

0

Notodontidae

Mimopydna pallida (Butler, 1877)

herb

specialist

1

0

0

0

Notodontidae

Nerice bipartita Butler, 1885

woody

specialist

0

1

0

0

Notodontidae

Peridea gigantea Butler, 1877

woody

specialist

1

0

0

0

Notodontidae

Phalera flavescens (Bremer & Grey, 1853)

woody

specialist

1

0

0

0

Notodontidae

Phalerodonta manleyi (Leech, 1889)

woody

specialist

0

1

0

0

Notodontidae

Pheosiopsis cinerea (Butler, 1879)

woody

specialist

1

0

0

0

Notodontidae

Pterostoma gigantinum Staudinger, 1892

woody

specialist

1

0

0

0

Notodontidae

Shaka atrovittatus (Bremer, 1861)

woody

specialist

0

1

1

0

Notodontidae

Stauropus basalis (Moore, 1877)

woody

generalist

1

0

0

1

Notodontidae

Stauropus fagi (Linnaeus, 1758)

woody

generalist

1

0

0

0

Notodontidae

Syntypistis cyanea (Leech, 1889)

woody

specialist

1

0

0

0

Oecophoridae

Acryptolechia malacobyrsa (Meyrick, 1921)

dead leaves

generalist

0

0

0

1

Oecophoridae

Acryptolechia sp. 1

unknown

unknown

0

1

1

0

Oecophoridae

Periacma delegata Meyrick, 1914

unknown

unknown

0

1

1

0

Oecophoridae

Tyrolimnas anthraconesa Meyrick, 1934

dead leaves

generalist

0

1

0

0

Pyralidae

Acrobasis bellulella (Ragonot, 1893)

woody

generalist

1

0

0

0

Pyralidae

Acrobasis birgitella (Roesler, 1975)

woody

specialist

1

0

0

0

Pyralidae

Acrobasis ferruginella Wileman, 1911

woody

specialist

1

0

0

0

Pyralidae

Acrobasis squalidella Christoph, 1881

woody

specialist

0

1

0

1

Pyralidae

Addyme confusalis Yamanaka, 2006

herb/woody

generalist

1

1

0

1

Pyralidae

Assara funerella (Ragonot, 1901)

woody

specialist

1

0

0

0

Pyralidae

Ceroprepes ophthalmicella (Christoph, 1881)

woody

specialist

1

0

0

0

Pyralidae

Ceroprepes patriciella Zeller, 1867

unknown

unknown

1

0

0

0

Pyralidae

Ectomyelois pyrivorella (Matsumura, 1899)

woody

specialist

1

0

0

0

Pyralidae

Emmalocera venosella (Wileman, 1911)

unknown

unknown

1

1

0

1

Pyralidae

Endotricha consocia (Butler, 1879)

unknown

unknown

0

1

0

1

Pyralidae

Endotricha kuznetzovi Whalley, 1963

dead leaves

generalist

1

1

1

0

Pyralidae

Endotricha olivacealis (Bremer, 1864)

dead leaves

generalist

1

1

0

1

Pyralidae

Etielloides bipartitellus (Leech, 1889)

unknown

unknown

1

0

0

0

Pyralidae

Euzophera batangensis Caradja, 1939

woody

generalist

1

0

0

0

Pyralidae

Furcata dichromella (Ragonot, 1893)

woody

specialist

1

0

0

0

Pyralidae

Furcata hollandella (Ragonot, 1893)

woody

specialist

1

0

0

0

Pyralidae

Furcata pseudodichromella (Yamanaka, 1980)

woody

specialist

1

0

0

0

Pyralidae

Hypsopygia regina (Butler, 1879)

woody

generalist

0

1

0

1

Pyralidae

Lepidogma melanobasis Hampson, 1906

unknown

unknown

0

1

0

1

Pyralidae

Locastra muscosalis (Walker, 1866)

woody

generalist

1

0

0

0

Pyralidae

Maliarpha borealis Sasaki, 2012

unknown

unknown

0

1

0

0

Pyralidae

Nyctegretis trigangulella (Hampson, 1901)

unknown

unknown

1

0

0

0

Pyralidae

Oncocera bitinctella (Wileman, 1911)

unknown

unknown

1

0

0

0

Pyralidae

Oncocera semirubella (Scopoli, 1763)

herb

specialist

1

1

1

1

Pyralidae

Orthaga achatina (Butler, 1878)

woody

generalist

0

0

0

1

Pyralidae

Orthaga euadrusalis Walker, 1859

woody

specialist

0

0

0

1

Pyralidae

Ortholepis infausta (Ragonot, 1893)

unknown

unknown

1

0

0

0

Pyralidae

Orthopygia placens (Butler, 1879)

unknown

unknown

1

0

0

0

Pyralidae

Paraemmalocera gensanalis (South, 1901)

unknown

unknown

1

0

0

0

Pyralidae

Patagoniodes nipponellus (Ragonot, 1901)

unknown

unknown

1

0

0

0

Pyralidae

Phycitodes matsumurellus (Shibuya, 1927)

unknown

unknown

0

1

0

0

Pyralidae

Pseudacrobasis nankingella Roesler, 1975

unknown

unknown

0

1

0

0

Pyralidae

Quasipuer colon (Christoph, 1881)

unknown

unknown

1

0

0

1

Pyralidae

Sacada fasciata (Butler, 1878)

woody

generalist

1

0

0

1

Pyralidae

Salma amica (Butler, 1879)

unknown

unknown

1

0

0

0

Pyralidae

Salma elegans (Butler, 1881)

herb/woody

generalist

1

1

0

0

Pyralidae

Scirpophaga virginia Schultze, 1908

unknown

unknown

1

0

0

0

Pyralidae

Scirpophaga xanthopygata Schawerda, 1922

unknown

unknown

1

0

0

0

Pyralidae

Selagia spadicella (Hübner, 1796)

unknown

unknown

0

0

0

1

Pyralidae

Tegulifera bicoloralis (Leech, 1889)

dead leaves

generalist

1

0

0

0

Pyralidae

Termioptycha margarita (Butler, 1879)

unknown

unknown

1

0

0

1

Pyralidae

Trebania flavifrontalis (Leech, 1889)

unknown

unknown

0

1

0

0

Saturniidae

Actias aliena (Butler, 1879)

woody

generalist

1

0

0

0

Saturniidae

Saturnia japonica (Moore, 1872)

woody

generalist

1

0

0

0

Saturniidae

Saturnia jonasii (Butler, 1877)

woody

generalist

0

1

1

1

Sphingidae

Acosmeryx naga (Moore, 1858)

woody

specialist

1

1

0

1

Sphingidae

Agrius convolvuli (Linnaeus, 1758)

herb

generalist

0

0

1

0

Sphingidae

Ambulyx ochracea Butler, 1885

woody

specialist

1

0

0

0

Sphingidae

Ampelophaga rubiginosa Bremer & Grey, 1853

woody

specialist

1

1

0

1

Sphingidae

Callambulyx tatarinovii (Bremer & Grey, 1852)

woody

specialist

1

0

0

0

Sphingidae

Clanis bilineata (Walker, 1866)

herb/woody

generalist

1

1

0

0

Sphingidae

Deilephila askoldensis (Oberthür, 1879)

herb

specialist

1

1

1

0

Sphingidae

Deilephila elpenor (Linnaeus, 1758)

herb

generalist

1

0

0

0

Sphingidae

Marumba gaschkewitschii (Bremer & Grey, 1853)

herb/woody

specialist

1

0

1

1

Sphingidae

Marumba sperchius (Ménétriès, 1857)

woody

specialist

1

0

0

0

Sphingidae

Smerinthus planus Walker, 1856

woody

generalist

1

0

0

0

Sphingidae

Theretra japonica (Boisduval, 1869)

herb/woody

generalist

1

0

1

1

Thyrididae

Striglina cancellata (Christoph, 1881)

woody

specialist

1

1

0

0

Tortricidae

Aethes cnicana (Westwood, 1854)

unknown

unknown

1

0

0

0

Tortricidae

Aethes rectilineana (Caradja, 1939)

unknown

unknown

1

0

0

0

Tortricidae

Ancylis mandarinana Walsingham, 1900

woody

specialist

1

0

0

0

Tortricidae

Apotomis basipunctana (Walsingham, 1900)

woody

specialist

0

0

0

1

Tortricidae

Apotomis capreana (Hübner, 1817)

woody

generalist

1

0

0

0

Tortricidae

Apotomis geminata (Walsingham, 1900)

woody

generalist

1

0

0

0

Tortricidae

Archips audax Razowski, 1977

herb/woody

generalist

1

0

0

0

Tortricidae

Archips fuscocupreana Walsingham, 1900

herb/woody

generalist

1

0

0

0

Tortricidae

Archips ingentana (Christoph, 1881)

herb/woody

generalist

1

0

0

0

Tortricidae

Archips oporana (Linnaeus, 1758)

woody

generalist

1

0

0

0

Tortricidae

Archips semistructa (Meyrick, 1937)

herb/woody

generalist

1

0

0

0

Tortricidae

Choristoneura longicellana (Walsingham, 1900)

woody

generalist

1

0

0

0

Tortricidae

Cochylidia subroseana (Haworth, 1811)

herb

specialist

1

0

0

0

Tortricidae

Cydia danilevskyi (Kuznetzov, 1973)

woody

specialist

1

0

0

0

Tortricidae

Diplocalyptis congruentana (Kennel, 1901)

herb

specialist

0

0

0

1

Tortricidae

Epiblema autolitha (Meyrick, 1931)

unknown

unknown

1

0

0

0

Tortricidae

Epiblema foenella (Linnaeus, 1758)

herb

specialist

1

1

1

1

Tortricidae

Epinotia bicolor (Walsingham, 1900)

woody

specialist

1

0

0

0

Tortricidae

Epinotia majorana (Caradja, 1916)

herb

specialist

1

0

0

0

Tortricidae

Eucosma catharaspis (Meyrick, 1922)

unknown

unknown

1

0

0

0

Tortricidae

Eucosma denigratana (Kennel, 1901)

unknown

unknown

1

0

0

0

Tortricidae

Eucosma metzneriana (Treitschke, 1830)

herb

specialist

1

0

0

0

Tortricidae

Eucosma striatiradix Kuznetzov, 1964

unknown

unknown

1

0

0

0

Tortricidae

Eucosma tundrana (Kennel, 1900)

unknown

unknown

0

1

0

0

Tortricidae

Eudemis porphyrana (Hübner, 1799)

woody

specialist

0

1

0

0

Tortricidae

Eugnosta dives (Butler, 1878)

herb

specialist

0

0

1

0

Tortricidae

Eupoecilia ambiguella (Hübner, 1976)

woody

generalist

0

0

0

1

Tortricidae

Eupoecilia inouei Kawabe, 1972

unknown

unknown

1

1

0

0

Tortricidae

Gravitarmata margarotana (Heinemann, 1863)

woody

specialist

1

0

0

0

Tortricidae

Gynnidomorpha vectisana (Humphreys & Westwood, 1845)

herb

specialist

1

0

0

0

Tortricidae

Gypsonoma dealbana (Frölich, 1828)

woody

generalist

0

0

0

1

Tortricidae

Hedya auricristana (Walsingham, 1900)

woody

generalist

1

0

0

0

Tortricidae

Lobesia reliquana (Hübner, 1776)

unknown

unknown

1

0

0

0

Tortricidae

Matsumuraeses phaseoli (Matsumura, 1900)

herb

specialist

1

0

0

0

Tortricidae

Neoanathamna nipponica (Kawabe, 1976)

dead leaves

generalist

0

0

0

1

Tortricidae

Neocalyptis angustilineata (Walsingham, 1900)

dead leaves

generalist

1

0

0

0

Tortricidae

Olethreutes examinata (Falkovitsh, 1966)

unknown

unknown

1

0

0

0

Tortricidae

Pandemis dumetana (Treitschke, 1835)

herb/woody

generalist

1

1

1

1

Tortricidae

Phaecasiophora roseana (Walsingham, 1900)

woody

specialist

1

1

0

1

Tortricidae

Phtheochroa pistrinana (Erschoff, 1877)

herb

specialist

1

0

0

1

Tortricidae

Ptycholoma lecheana (Linnaeus, 1758)

woody

generalist

1

0

0

0

Tortricidae

Saliciphaga acharis (Butler, 1879)

woody

specialist

1

0

0

0

In order to explore the food habits of moth species at each site in both years, the moth species were classified into six feeding groups: 1) species that primarily feed on herbs, 2) species that primarily feed on shrubs and/or trees, 3) species that feed on both herbs and trees, 4) species that primarily feed on lichens, 5) species that primarily feed on dead leaves and 6) species whose food habit is unknown, referring to the guide “The Standard of Moths in Japan I-IV” (Kishida 2011a, Kishida 2011b, Hirowatari et al. 2013, Nasu et al. 2013). The collected moths were also classified into two groups based on the range of host plants: 1) specialist, defined as species that feed on only one plant family and 2) generalists, defined as species that feed on more than two plant families. We calculated the proportions of each feeding groups for each site in both years.

Results and Discussion

The number of taxa of moths recorded in 2018 at each site (excluding the 4 families and 10 species of micro-moths) was 14 families, 79 genera and 95 species for the grassland site; 16 families, 115 genera and 137 species for the forest site and 14 families, 103 genera and 123 species for the mixed site (Fig. 2). In total, 17 families, 181 genera and 226 species were recorded in 2018. This was nearly two-thirds of the species recorded in 1987, which comprised of 16 families, 291 genera and 396 species. Amongst all the moth species documented, only 118 species were observed in both periods. The 278 species that were recorded in 1987 were not recorded in 2018, while 115 species were newly recorded in 2018.

Figure 2.  

Number of moths collected in each site in both years in each month.

Species composition of moths between years and amongst sites

The results of the PERMANOVA showed that the species composition of the moths significantly differed between years (Fig. 3a; R2 = 0.06, P = 0.02). This suggests that species composition at the three sites in 2018 differed from that in 1987. On the other hand, when the data of 1987 was excluded from the analysis, it did not differ amongst sites in 2018 (Fig. 3b; R2 = 0.08, P = 0.99). This may be because the three sites in 2018 were located in a geographically adjacent area (an average of 400 m between sites). Studies that detected differences in moth species composition amongst different vegetation and management were sites that were geographically separated by more than 1 km (Öckinger and Smith 2006, Pöyry et al. 2009). As some moth species are known to show high mobility (i.e. a flight range larger than 500 m in radius) (van der Meulen and Groenendijk 2005, Merckx and Macdonald 2015), species composition of moths may not differ in a small spatial scale. Furthermore, because the present data included only one replicate for each type of vegetation, more replicates from multiple sites would be useful in confirming that the difference in vegetation reflecting the management did not affect the moth species composition in the semi-natural grassland in Kanpu-zan.

Figure 3.  

Non-metric multidimensional scaling (NMDS) ordination graphs of monthly aggregated moth communities showing (a) for both years and (b) for 2018.

Food habits

The number of species that may rely on woody plants comprised more than half of the total number of species at all sites in both years (Fig. 4a). This includes species that feed only on woody plants; species that feed on both herbs and woody plants; species that feed on lichens and mosses; and species that feed on dead leaves. These species were known to consume woody plants that belong to the families of Fagaceae, Rosaceae, Fabaceae and Caprifoliaceae. Trees and shrubs that belong to Fagaceae and Rosaceae were dominant in the forest site and Fabaceae shrubs and vines were scattered across the grassland site, while the mixed site had a mixture of vegetation of both sites. Moreover, trees and shrubs such as Cerasus jamasakura H.Ohba and Quercus serrata Murray were present in grasslands in 1987 (Takahashi 1993, K. Umetsu, personal observation). These suggest that, in both years, moth fauna in Kanpu-zan was mainly comprised of species that may feed on woody plants.

Figure 4.  

Proportion of each food habits for moth species (a) of each site in both years, (b) those unique to each year and common in both years.

For the 118 species that were recorded in both in 1987 and 2018, the composition rate of species that may use both herbs and woody plants was 19%, which was higher compared to that for the species that were only recorded in either 1987 or 2018, at 10.1% and 11.2%, respectively (Fig. 4b). This may be because the species that can feed on both herbs and woody species are adapted to habitats with various vegetations and were tolerant to the environmental change between 1987 and 2018 caused by decline in grasslands. Species that primarily feed on herbs such as Deilephila askoldensis, which is known to inhabit volcanic grasslands (Yano 2011), consisted of 15-20% in all three sites in 2018. The semi-natural grasslands in Kanpu-zan may be a valuable habitat for the larvae of these moths. On the other hand, the composition rate of species that feed on lichens was relatively low in all sites and especially low at the grassland site in 2018 (Fig. 4a). This is consistent with the result in 1987, which identified that it is one of the characteristics in grasslands that lichen feeding moths are scarce (Takahashi 1993).

The composition rate of specialist and generalist moths was stable across sites including the year 1987. Generalists consisted of 43.8±1.5% (mean±SE) of the total number of moth species in both years (Fig. 5). Out of the 280 species of moths that were only recorded in 1987, 107 species were generalists that feed on plants that are commonly distributed in Kanpu-zan (Masui et al. 2017).

Figure 5.  

Proportion of food range (specialist/generalist) for moth species (a) of each site in both years, (b) those unique to each year and common in both years.

Conclusion

The present study suggests the species composition changed between 1987 and 2018 and the number of moth species largely declined in the last 30 years. Wenzel et al. (2006) also showed that the number of butterflies and burnet moths had declined between 1972 and 2001 in calcareous grassland remnants in south-western Germany. They suggested that species which require structured habitats, species with low mobility, species which require more than 16 ha of habitat and specialist feeders were especially affected by the decline. As we did not find any moths that depend on endangered plants or whose food plant had become extinct, it is unlikely that the decline in the moths in Kanpu-zan was due to the loss in food plants. Other factors such as plant height, flowering species richness and habitat connectivity (Pöyry et al. 2009) may have caused the decline. Continuing landscape management such as mowing and burning may be important, not only for endangered species, but also for common moth species and to maintain the nocturnal moth community in semi-natural grasslands.

A previous study in Japan covering 31 agricultural areas, including semi-natural grasslands, demonstrated that declines in herbivorous insects in both abandoned and intensified use of agricultural landscapes are also explained by multiple factors (Uchida and Ushimaru 2014). It has been reported that there are large differences with respect to the preferred management intensity amongst species groups in the grassland habitats (Pöyry et al. 2006). More studies on other organism fauna are needed for understanding the conservation of semi-natural grassland, considering that the loss of semi-natural grasslands is one of the major threats to biodiversity.

Acknowledgements

We are grateful to K. Umetsu for help on the species identification and advice on the study. We thank to M. Takahashi for advice on the fieldwork. We appreciate S. Tsuda for helpful advice on the field site. We also thank D. Plotkin and J. Awad and an anonymous reviewer for their constructive comments on our manuscript.

References

Supplementary material

Suppl. material 1: monthly species occurences 
Authors:  Masaru Kamikura, Yuzu Sakata
Data type:  occurrences
Brief description: 

monthly species occurrences of nocturnal moths