Fauna of nocturnal moth species collected in a semi-natural grassland at Kanpu-zan in northern Japan.

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.

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 , 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. , , ) 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 , , and 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.

Table 1.

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 , , and . Trees and shrubs that belong to and were dominant in the forest site and 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 H.Ohba and 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 , 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.

monthly species occurences

Data type: occurrences

Brief description: monthly species occurrences of nocturnal moths

File: oo_320380.csv