Diversity and Distribution Patterns of Geometrid Moths (Geometridae, Lepidoptera) in Mongolia

Abstract

Geometrids are a species-rich group of moths that serve as reliable indicators for environmental changes. Little is known about the Mongolian moth fauna, and there is no comprehensive review of species richness, diversity, and distribution patterns of geometrid moths in the country. Our study aims to review the existing knowledge on geometrid moths in Mongolia. We compiled geometrid moth records from published scientific papers, our own research, and from the Global Biodiversity Information Facility (GBIF) to produce a checklist of geometrid moths of Mongolia. Additionally, we analyzed spatial patterns, species richness, and diversity of geometrid moths within 14 ecoregions of Mongolia and evaluated environmental variables for their distribution. In total, we compiled 1973-point records of 388 geometrid species. The most species-rich ecoregion in Mongolia was Daurian Forest Steppe with 142 species. Annual precipitation and maximum temperature of the warmest month were the most important environmental variables that correlated with NMDS axes in an analysis of geometrid assemblages of different ecoregions in Mongolia.

1. Introduction

Regarded as disturbing pests or less charismatic than butterflies, moths are nevertheless creatures with an important role in the ecosystem and the potential to serve as environmental indicators [1,2,3,4]. Moths are globally distributed and it is estimated that more than 130,000 described species exist [5], far more than the more conspicuous and mostly diurnal butterflies with ca. 20,000 species. Many moths are pollinators, but due to their nocturnal activity they are not well studied [6]. In a recent review from the current literature, Hahn and Brühl reported that in Europe and North America there are 227 moth–plant interactions with 129 moth species involved [6]. Geometrid moths (Geometridae), constituting one of the biggest families of Lepidoptera, are a species-rich and easily recognizable family that have served as indicators for environmental changes in many previous studies [7,8,9,10]. These groups also appear to be effective at colonizing habitats after natural or anthropogenic disturbances [11]. There are approximately 24,000 described species of Geometridae worldwide [12]. Although Mongolia is one of the largest countries (rank 19th in size) on Earth, little is known about its moth fauna, and there is no comprehensive review of species richness, diversity, and distribution patterns of geometrid moths in the country. A few researchers attempted to summarize information to mainly confirm this lack of information [13].

Mongolia is a country that encompasses landscapes with a high variety of climatic and geographic features with forest in the north, high mountains in the west, desert in the south, and steppes in the eastern and central parts of Mongolia [14,15]. Altogether, it comprises 16 ecoregions [16] (Figure 1). Ecosystems change along a latitudinal gradient from forest in the north, over steppe and semi-desert to desert in the south [17]. In most areas of the country, livestock herding is a dominant land-use practice, and due to overgrazing, some pasture lands have recently been degraded [18]. With recent discoveries of various mineral resources, mining has become not only the main economic sector, but also the major reason for environmental disturbance in Mongolia. Together with climate change, it is the major driver for habitat loss and environmental changes [14,19]. As a result of these anthropogenic changes, many species are disappearing, but there is little information about which species are at greatest risk of becoming extinct, especially for the less studied taxa.

In order to monitor diversity loss and gain, and to further study the influence of environmental disturbance and climate change on geometrid moths in Mongolia, we need an up-to-date dataset that mirrors the current state of knowledge and that includes all species already recorded. Given this knowledge gap, this study aims to review, summarize, and evaluate the existing knowledge on geometrid moths in Mongolia. It will provide a baseline for further studies, as well as define research priorities in the field. In this study, we aim to: (1) provide a checklist of geometrid moths of Mongolia, setting a baseline for future studies, (2) analyze distribution patterns and species richness and diversity of geometrid moths within ecoregions of Mongolia, and (3) analyze which environmental variables are most important in determining their distribution. We are aware that all results can only give a provisional status due to the data situation, especially the results for Objectives 2 and 3 can only be given with caution; however, our detailed review of the current data will help to define the needs for further research more efficiently.

Study Review

Information on the species composition of Macrolepidoptera of Mongolia began to accumulate from the end of the nineteenth century, as a result of the works of collectors such as Fritz Dörries, Hauberhauer and Leder, and others. Otto Staudinger [20] published the first paper on the collection of Fritz Dörries, who made a trip in 1879 to Khentii Mountains to collect Lepidoptera. This resulted in data on the location of 75 species of geometrids in central and western parts of Mongolia [20]. Later, Staudinger published several papers and books on the fauna of Palaearctic Lepidoptera which included some geometrid species from Mongolia [21,22,23]. In 1964, a Mongolian–German expedition conducted a biological survey, as a result of the expedition 214 Lepidopteran exemplars were sampled. Burchard Alberti later published the results on Lepidoptera and nine geometrid species were listed in the paper [24]. Likewise, Joseph Moucha listed four geometrid species from a Mongolian–Czech entomological–botanical expedition, which was conducted around 1960 [25]. Grigory Grum-Grshimailo found three geometrid species from Selenge Aimag in the collection of M.I. Molleson [26]. Alexander Mikhailovich Djakonov [27,28] recorded a new occurrence of Horisme scosiata and described one new species Scotopteryx transbaicalica from the family of Geometridae based on old material of Staudinger. Other researchers such as Karl Dietze [29], Eugen Wehrli [30], and Fritz Heydemann [31] also described new species. In the fourth volume and its supplementary of “Die Gross-Schmetterlinge der Erde. Die Spanner des Palaearktischen Faunengebietes“ series edited by Adalbert Seitz, 34 geometrid species were listed for Mongolia [32,33].

The most important contribution to the collection and study of Mongolian geometrid moths were made by Russian and Soviet expeditions led by Pyotr Kuzmich Kozlov and later by Soviet–Mongolian expeditions [34,35,36]. During the survey of Soviet–Mongolian expeditions, Jaan Viidalepp recorded a total of 201 geometrid species.Viidalepp later in 1999 compiled a checklist of geometrid moths of the former U.S.S.R and in this monograph 210 species were included for Mongolia [37]. Particularly rich and diverse material on Lepidoptera (41,000 specimens) were collected by the Hungarian expeditions conducted by Zoltán Kaszab, who made six entomological collecting trips along latitudinal and longitudinal gradients in Mongolia, between 1963 and 1968. András Vojnits published several papers based on the Kaszab’s collections dedicated to subfamilies of Geometridae in the period between 1974 and 1979. He recorded 177 species from the whole collection, described 39 species new to the fauna of Mongolia and four species new to science [38,39,40,41,42,43,44]. Malcolm J. Scoble [45] presented 66 taxa from Mongolia.

Other researchers also contributed to the study of Mongolian geometrid moths. For instance, Gantigmaa Ch. and coworkers recorded 90 species in the West Khentii of Northern Mongolia [46]. In the book “Biodiversity of Sokhondinsky Reserve”, 29 geometrid species from Mongolia have been included [47]. Beljaev and Vasilenko [48] noted 29 species of geometrid moths in Mongolia. Vasilenko and colleagues [49,50,51] recorded eight species and described one new species Rhodostrophia ustyuzhanini in Western Mongolia. In 2012 and 2013, we collected 70 geometrid species from central and northern parts of Mongolia [4]. Mironov and Glasworthy [52] reported 57 species with two species (Eupithecia ankini, Eupithecia munguata) new to science and 12 species new to the fauna of Mongolia. Erlacher et al. studied six geometrid species from Mongolia and described one new species Charissa beljaevi [53,54,55]. In 2019, Makhov and Beljaev [56] studied the geometrid moths of the Baikal Region and recorded 14 species from Mongolia. In six volumes of “The Geometrid Moths of Europe”, 117 moth species are listed from Mongolia. We validated our species checklist with these volumes [57,58,59,60,61,62].

2. Materials and Methods

We compiled geometrid moth records from published scientific papers, from our work [63] (all sample identifications were double checked by curator T. Enkhbayar, Department of Biology, National University of Mongolia), from the collections of the Siberian Zoological Museum (curator - S.V.Vasilenko) [64], and also from the Global Biodiversity Information Facility (GBIF) [65]. Lastly, we checked the “Revised, annotated systematic checklist of the Geometridae of Europe and adjacent areas, Vols 1–6” [62]. From the Museum collections we could only get country-level information, not the exact location. From GBIF data, we included 380 records into our species list [65]. Fourteen specimens of six species were found in the public data of The Barcode of Life Data System (Bold System) [66].

We used Google Scholar to search the literature with following search strings:

-

With all of the words: Mongol (in English Mongolia, in German Mongolei, thus it was better to use only Mongol);

-

With at least one of the words: Geometrid OR Larentiinae OR Desmobathrinae OR Ennominae OR Archiearinae OR Geometrinae OR Oenochrominae OR Orthostixinae OR Sterrhinae;

As a result of the search, 184 literatures appeared, though many of them were about geometrid moths of Inner Mongolia. These we excluded from our list.

- Without the words: Inner Mongolia.

After excluding Inner Mongolia, 96 results remained and of these, 73 were relevant to our study.

Totally, we compiled 1973-point records of 388 geometrid species (Table S1). Of these records, 87 species were missing information on exact locations, these 87 species are used to estimate species richness and listed in the species checklist but are excluded from other analysis. We georeferenced species locations from literature and generated coordinates of each location with Google Earth [67]. After that we cross-checked each species name in “The Global Lepidoptera Names Index” [68]. Moreover, experts on geometrid moths such as Axel Hausmann, Jaan Viidalepp, Gunnar Brehm, Sven Erlacher, and Pasi Sihvonen validated most species of our checklist and provided further literatures.

In the next step we used the sampled data in order to estimate true species richness, to evaluate the distribution of species within Mongolia, and to identify regions that have been undersampled so far by species rarefaction. For these reasons, we transformed all species locations into 2° × 2° grid cells, resulting in 51 grid cells inhabited by 301 species. Of 301 species, 121 were unique species occuring only once within 51 grids. To estimate species richness we applied Good Turing Theory, which uses unique species for estimation [69]. We used the application SuperDuplicates (https://chao.shinyapps.io/SuperDuplicates/) for the estimation with the following setting: Data type: incidence data; Number of observed species (SOBs): 388; Number of uniques (Q1): 208 (we combined the 121 unique species with the former mentioned 87 species without locations).

Further we calculated rarefaction curves for single ecoregions to assess collection quality in different areas of Mongolia. Four ecoregions (Altai Alpine Meadow and Tundra, Dzungarian Basin Semi-Desert, Khangai Mountains Alpine Meadow and Sayan Alpine Meadows, and Tundra) were strongly under sampled, having species richness below 15, thus we excluded them from the analysis to avoid misleading interpretation.

To estimate the rarefaction curve across grid cells and ecoregions, we calculated interpolation and extrapolation of species richness using the ‘iNEXT’ package: Interpolation and extrapolation for species richness in R [70,71] with 0.95 confidence interval and prepared the rarefaction plots with ‘devtools’ package [72] and ggiNEXT function of ‘ggplot2′ package [73].

We performed Non-Metric Multidimensional Scaling Analysis (NMDS) to check the dissimilarity of geometrid species composition between ecoregions based on the zero-adjusted Bray–Curtis dissimilarity measure using ‘phytomosaic/ecole’ and ‘vegan’ package [74,75,76]. For estimation of pairwise similarities between ecoregions, we calculated the estimated abundance based Soerensen Index by abundance data using online program SpadeR [77]. We preferred Soerensen Index over Jaccard Index, while the result was a little bit higher than Jaccard. This estimated abundance based index can detect unseen shared species and is appropriate to evaluate beta diversity of samples under sampling bias [78].

We used 19 Bioclim data with 30 arc seconds resolution as climatic variables for the region [79]. We extracted these variables for the fourteen ecoregions. Ecoregion GIS data for Mongolia were downloaded from The Nature Conservancy (TNC) [80]. In two ecoregions no geometrid moths were found, namely, Khangai Mountains Conifer Forests and Sayan Intermontane Steppe (Figure 1). We thus excluded these ecoregions from the further analysis. To check for strong linear dependencies among explanatory variables we computed the variance inflation factor (VIF) for each variable in R package ‘vegan’. We excluded variables with VIF values higher than 10 [81] (

Table 1. List of the environmental [79] variables* for the fourteen ecoregions used in this study. All variables have been entered into forward selection method for selecting most important variables. The selected variables were later fitted in the Non-Metric Multidimensional Scaling Analysis (NMDS). Colors refer to the map in Figure 1.

Ecoregions	Bio1	Bio2	Bio5	Bio6	Bio7	Bio10	Bio11	Bio12	Biome [83]
Alashan Plateau Semi-Desert	5.1	14.1	28.6	−20.3	49	20.6	−11.7	85	Deserts and Xeric Shrublands
Altai Alpine Meadow and Tundra	−4.5	12.3	17.1	−28.1	45.2	10.3	−20.3	199	Montane Grasslands and Shrublands
Altai Montane Forest and Forest Steppe	−1.8	13.1	20.5	−26.8	47.3	13.4	−18.5	148	Temperate Conifer Forests
Dzungarian Basin Semi-Desert	3.9	14	27.4	−23	50.4	19.6	−13.9	91	Deserts and Xeric Shrublands
Daurian Forest Steppe	−1.5	13.9	23.7	−29.1	52.9	16	−21	306	Temperate Grasslands, Savannas and Shrublands
Eastern Gobi Desert Steppe	3.3	13.4	27.6	−22.5	50.1	19.8	−14.7	130	Deserts and Xeric Shrublands
Gobi Lakes Valley Desert Steppe	0.7	14.6	23.8	−24.3	48.1	15.9	−15.5	141	Deserts and Xeric Shrublands
Great Lakes Basin Desert Steppe	−1.6	13.5	24.2	−31.7	55.9	16.6	−23.1	147	Deserts and Xeric Shrublands
Khangai Mountains Alpine Meadow	−5.6	14.3	17.3	−30.5	47.8	9.7	−22.1	261	Montane Grasslands and Shrublands
Mongolian-Manchurian Grassland	0.3	13.6	25.4	−26.4	51.8	17.6	−18.7	224	Temperate Grasslands, Savannas and Shrublands
Sayan Alpine Meadows and Tundra	−8.4	13.6	16.3	−34.9	51.2	8.5	−27.3	355	Montane Grasslands and Shrublands
Sayan Montane Coniferous Forests	−5.1	13.7	19.2	−31.3	50.4	11.4	−23.5	381	Temperate Conifer Forests
Selenge-Orkhon Forest Steppe	−3.2	14.3	20.6	−29.7	50.3	12.9	−21.4	277	Temperate Grasslands, Savannas and Shrublands
Trans-Baikal Coniferous Forests	−3.3	13.4	22.1	−31.1	53.2	14.6	−23.3	366	Boreal Forests/Taiga

*Environmental variables with VIF under 10. Bio1—Annual Mean Temperature [°C]; Bio2—Mean Diurnal Range [°C]; Bio5—Max Temperature [°C]; Bio6 - Min Temperature [°C]; Bio7—Temperature Annual Range [°C]; Bio10—Mean Temperature of Warmest Quarter [°C]; Bio11—Mean Temperature of Coldest Quarter [°C]; Bio12—Annual precipitation [mm].

). We chose the most significant environmental variables with forward selection method by using vegan’s ‘ordistep’ function [81]. Variables selected by forward selection method were fitted into the ordination plot using vegan’s ‘entfit’ function.

All analysis were performed in R [82] and most graphs were made with package ‘ggplot2′ [73].

3. Results

Altogether, we recorded 388 geometrid species of six subfamilies: Archiearinae, Desmobathrinae, Ennominae, Geometrinae, Larentiinae, and Sterrhinae (Appendix A

Table A1. Checklist of geometrid moths in Mongolia. Note that we conducted all analysis at species level. Here subspecies are listed to show compiled data in more detail. The listed references include in most cases articles with location information.

Subfamily	Species	Author	Year	Reference
Archiearinae	Archiearis notha	Hübner	1802	[34]
Archiearinae	Archiearis parthenias	Linnaeus	1761	[34]
Archiearinae	Archiearis parthenias sajana	Prout	1912	[46]
Archiearinae	Leucobrephos middendorfii	Ménétriés	1858	[41]
Desmobathrinae	Gypsochroa renitidata	Hübner	1817	[57]
Ennominae	Abraxas grossulariata	Linnaeus	1758	[21,34,46,63,65]
Ennominae	Abraxas grossulariata dsungarica	Wehrli	1939	[38]
Ennominae	Alcis deversata	Staudinger	1892	[34,39,46,63,65]
Ennominae	Alcis extinctaria	Eversmann	1851	[23,34,36,39,65,91]
Ennominae	Alcis jubata	Thunberg	1788	[37]
Ennominae	Alcis repandata	Linnaeus	1758	[65]
Ennominae	Alloharpina conjungens	Alphéraky	1892	[33]
Ennominae	Amraica superans	Butler	1878	[33]
Ennominae	Angerona prunaria	Linnaeus	1758	[24,34,46,63,65]
Ennominae	Angerona prunaria kentearia	Staudinger	1892	[39]
Ennominae	Angerona prunaria mongoligena	Bryk	1949	[62]
Ennominae	Apeira syringaria	Linnaeus	1758	[63]
Ennominae	Apocheima hispidaria	Denis & Schiffermüller	1775	[34]
Ennominae	Apocolotois almatensis	Djakonov	1952	[39]
Ennominae	Apocolotois smirnovi	Romanoff	1885	[39]
Ennominae	Arichanna barteli	Prout	1915	[32,45]
Ennominae	Arichanna melanaria	Linnaeus	1758	[34,46,65,91]
Ennominae	Arichanna melanaria decolorata	Staudinger	1892	[45]
Ennominae	Arichanna melanaria praeolivina	Wehrli	1933	[39]
Ennominae	Aspitates conspersaria	Staudinger	1901	[23,45]
Ennominae	Aspitates curvaria	Eversmann	1852	[1,8,14]
Ennominae	Aspitates forbesi	Munroe	1963	[65]
Ennominae	Aspitates gilvaria	Denis & Schiffermüller	1775	[23,24,34,36,63,91]
Ennominae	Aspitates gilvaria minimus	Vojnits	1975	[39]
Ennominae	Aspitates insignis	Alphéraky	1883	[36,39]
Ennominae	Aspitates kozhantchikovi	Munroe	1963	[36,65]
Ennominae	Aspitates mongolicus	Vojnits	1975	[39,65]
Ennominae	Aspitates mundataria	Stoll	1782	[34,46,63,65]
Ennominae	Aspitates mundataria uncinataria	Vojnits	1975	[39]
Ennominae	Aspitates obscurata	Wehrli	1953	[33,34,39]
Ennominae	Aspitates staudingeri	Vojnits	1975	[39]
Ennominae	Aspitates taylorae sibirica	Djakonov	1955	[36,65]
Ennominae	Aspitates tristrigaria	Bremer & Grey	1853	[34,37]
Ennominae	Astegania honesta	Prout	1908	[34]
Ennominae	Biston betularia	Linnaeus	1758	[34,46,63,91]
Ennominae	Biston betularia sibiricus	Fuchs	1899	[37]
Ennominae	Cabera exanthemata	Scopoli	1763	[23,34,46,65]
Ennominae	Cabera exanthemata hamica	Wehrli	1939	[39]
Ennominae	Cabera pusaria	Linnaeus	1758	[34,39,63]
Ennominae	Calcaritis pallida	Hedemann	1881	[47]
Ennominae	Chariaspilates formosaria	Eversmann	1837	[37]
Ennominae	Charissa agnitaria	Staudinger	1897	[55]
Ennominae	Charissa ambiguata	Duponchel	1830	[34,36,46,65]
Ennominae	Charissa ambiguata ophthalmicata	Lederer	1853	[39]
Ennominae	Charissa beljaevi	Erlacher et al., 2017	2017	[55]
Ennominae	Charissa bidentatus	Shchetkin & Viidalepp	1980	[46]
Ennominae	Charissa creperaria	Erschoff	1877	[34,55,65]
Ennominae	Charissa difficilis	Alphéraky	1883	[21,24,34,39,65]
Ennominae	Charissa gozmanyi	Vojnits	1975	[14]
Ennominae	Charissa macguffini	Smiles	1979	[65]
Ennominae	Charissa ochrofasciata	Staudinger	1895	[21,30,34,36,39,55,65]
Ennominae	Charissa remmi	Viidalepp	1988	[56,63]
Ennominae	Charissa sibiriata	Guenée	1858	[21,24,30,34,36]
Ennominae	Charissa subsplendidaria	Wehrli	1922	[63,92]
Ennominae	Charissa turfosaria	Wehrli	1922	[30,34,39,45,62]
Ennominae	Charissa vastaria	Staudinger	1892	[30,34]
Ennominae	Chiasmia aestimaria	Hübner	1809	[65]
Ennominae	Chiasmia aestimaria kuldschana	Wehrli	1940	[39]
Ennominae	Chiasmia clathrata	Linnaeus	1758	[23,24,26,34,36,46,63,65,91]
Ennominae	Chiasmia clathrata djakonovi	Kardakoff	1928	[38,39]
Ennominae	Chiasmia saburraria	Eversmann	1851	[21,34,65]
Ennominae	Chiasmia saburraria kenteata	Staudinger	1892	[38]
Ennominae	Cleora cinctaria	Denis & Schiffermüller	1775	[34,46,63]
Ennominae	Colotois pennaria	Linnaeus	1760	[46]
Ennominae	Deileptenia ribeata	Clerck	1759	[63]
Ennominae	Digrammia rippertaria	Duponchel	1830	[34]
Ennominae	Ectropis crepuscularia	Denis & Schiffermüller	1775	[34,46]
Ennominae	Eilicrinia orias	Wehrli	1933	[45]
Ennominae	Elophos banghaasi	Wehrli	1922	[30,34,45]
Ennominae	Ematurga atomaria	Linnaeus	1758	[23,24,34,36,46,65]
Ennominae	Ematurga atomaria krassnojarscensis	Fuchs	1899	[39]
Ennominae	Ennomos autumnaria	Werneburg	1859	[46]
Ennominae	Epione repandaria	Hufnagel	1767	[34]
Ennominae	Epione vespertaria	Linnaeus	1767	[34,39]
Ennominae	Epirranthis diversata	Denis & Schiffermüller	1775	[63]
Ennominae	Erannis jacobsoni	Djakonov	1926	[34,46,65]
Ennominae	Gnophopsodos ravistriolaria	Wehrli	1922	[36]
Ennominae	Gnophopsodos ravistriolaria ravistriolaria	Wehrli	1922	[55]
Ennominae	Gnophopsodos stemmataria	Eversmann	1848	[39]
Ennominae	Gnophopsodos tholeraria	Püngeler	1901	[50]
Ennominae	Gnophos bipartitus	Vojnits	1975	[39]
Ennominae	Gnophos rubefactaria	Püngeler	1902	[37]
Ennominae	Heliomata glarearia	Denis & Schiffermüller	1775	[46]
Ennominae	Hypomecis punctinalis	Scopoli	1763	[46]
Ennominae	Hypomecis roboraria	Denis & Schiffermüller	1775	[23,34,39,63]
Ennominae	Hypoxystis pluviaria	Fabricius	1787	[34,46,63]
Ennominae	Isturgia altaica	Vojnits	1978	[43]
Ennominae	Isturgia arenacearia	Denis & Schiffermüller	1775	[63,91]
Ennominae	Isturgia arenacearia mongolica	Vojnits	1974	[38]
Ennominae	Isturgia falsaria	Alphéraky	1892	[34]
Ennominae	Isturgia halituaria	Guenée	1858	[48]
Ennominae	Isturgia kaszabi	Vojnits	1974	[38]
Ennominae	Isturgia murinaria	Denis & Schiffermüller	1775	[34,36]
Ennominae	Isturgia murinaria uralica	Wehrli	1937	[63]
Ennominae	Jankowskia bituminaria	Lederer	1853	[65]
Ennominae	Jankowskia bituminaria raddensis	Wehrli	1941	[93]
Ennominae	Lomaspilis marginata	Linnaeus	1758	[23,34,46,65]
Ennominae	Lomaspilis opis amurensis	Hedemann	1881	[38]
Ennominae	Lomographa buraetica	Staudinger	1892	[34]
Ennominae	Lomographa temerata	Denis & Schiffermüller	1775	[46]
Ennominae	Lycia hirtaria	Clerck	1759	[63]
Ennominae	Lycia lapponaria	Boisduval	1840	[37]
Ennominae	Macaria alternata	Denis & Schiffermüller	1775	[34,46,91]
Ennominae	Macaria artesiaria	Denis & Schiffermüller	1775	[34,38]
Ennominae	Macaria brunneata	Thunberg	1784	[36,38,91]
Ennominae	Macaria circumflexaria	Eversmann	1848	[38,46,63,91]
Ennominae	Macaria costimaculata	Graeser	1888	[34]
Ennominae	Macaria latefasciata	Staudinger	1896	[21,34]
Ennominae	Macaria liturata	Clerck	1759	[65]
Ennominae	Macaria liturata pressaria	Christoph	1893	[37]
Ennominae	Macaria loricaria	Eversmann	1837	[36]
Ennominae	Macaria notata	Linnaeus	1758	[34,63]
Ennominae	Macaria notata kirina	Wehrli	1940	[38]
Ennominae	Macaria serenaria	Staudinger	1896	[21,34]
Ennominae	Macaria signaria	Hübner	1809	[38,46]
Ennominae	Macaria wauaria	Linnaeus	1758	[34,36]
Ennominae	Megalycinia strictaria	Lederer	1853	[21,34,39,46,63]
Ennominae	Megametopon piperatum	Alphéraky	1892	[34,39,65]
Ennominae	Narraga fasciolaria	Hufnagel	1767	[34,63]
Ennominae	Odontopera bidentata	Clerck	1759	[21,35,40,47,66]
Ennominae	Odontopera bidentata exsul	Tchetrerikov	1905	[36,39]
Ennominae	Odontopera bidentata rava	Vojnits	1975	[39,65]
Ennominae	Ourapteryx persica	Ménétriés	1832	[34]
Ennominae	Ourapteryx sambucaria	Linnaeus	1758	[63,65]
Ennominae	Perconia strigillaria	Hübner	1787	[46,63]
Ennominae	Petrophora kaszabi	Vojnits	1978	[43]
Ennominae	Phaselia narynaria	Oberthür	1913	[49]
Ennominae	Phaselia serrularia	Eversmann	1847	[65]
Ennominae	Phthonandria emaria	Bremer	1864	[39]
Ennominae	Plagodis dolabraria	Linnaeus	1767	[34]
Ennominae	Plagodis pulveraria	Linnaeus	1758	[21,34,65]
Ennominae	Plagodis pulveraria singularis	Vojnits	1975	[39]
Ennominae	Pleogynopteryx bituminaria	Lederer	1853	[21,34,39]
Ennominae	Pseudopanthera macularia	Linnaeus	1758	[34]
Ennominae	Pseudopanthera macularia cryptica	Beljaev	1997	[94]
Ennominae	Selenia dentaria	Fabricius	1775	[39]
Ennominae	Selenia dentaria alpestris	Wehrli	1940	[37]
Ennominae	Selenia ononica	Kostjuk	1991	[37]
Ennominae	Selenia sordidaria	Leech	1897	[39]
Ennominae	Selenia tetralunaria	Hufnagel	1767	[34,36,46,63]
Ennominae	Siona lineata	Scopoli	1763	[23,26,34,36,39,46,63,65]
Ennominae	Spartopteryx kindermannaria	Staudinger	1871	[36,39,46]
Ennominae	Xandrames dholaria	Moore	1868	[33]
Ennominae	Yezognophos vittaria	Thunberg	1792	[65]
Geometrinae	Chlorissa viridata	Linnaeus	1758	[34]
Geometrinae	Dyschloropsis impararia	Guenée	1858	[21,24,34,40,41,65]
Geometrinae	Geometra papilionaria	Linnaeus	1758	[40,46,63]
Geometrinae	Geometra papilionaria herbacearia	Ménétriés	1859	[41,65]
Geometrinae	Hemistola chrysoprasaria	Esper	1794	[46,63]
Geometrinae	Hemistola chrysoprasaria lissas	Prout	1912	[40]
Geometrinae	Hemistola zimmermanni	Hedemann	1879	[34,40,41]
Geometrinae	Hemithea aestivaria	Hübner	1799	[46]
Geometrinae	Jodis lactearia	Linnaeus	1758	[37]
Geometrinae	Microloxia herbaria	Hübner	1813	[34,65]
Geometrinae	Microloxia herbaria advolata	Eversmann	1837	[41]
Geometrinae	Thalera chlorosaria	Graeser	1890	[34,40,41,91]
Geometrinae	Thalera fimbrialis	Scopoli	1763	[63]
Geometrinae	Thetidia atyche	Prout	1935	[40,41]
Geometrinae	Thetidia chlorophyllaria	Hedemann	1879	[37]
Geometrinae	Thetidia correspondens	Alpheraky	1883	[49]
Geometrinae	Thetidia volgaria	Guenée	1858	[21,34,40,46,65]
Geometrinae	Thetidia volgaria mongolica	Staudinger	1897	[41]
Larentiinae	Acasis appensata	Eversmann	1842	[46,65]
Larentiinae	Anticlea badiata	Denis & Schiffermüller	1775	[34,63]
Larentiinae	Anticlea derivata	Denis & Schiffermüller	1775	[24,34,46,63]
Larentiinae	Aplocera plagiata roddi	Vasilenko	1995	[59]
Larentiinae	Baptria tibiale	Esper	1804	[34,42]
Larentiinae	Camptogramma bilineata	Linnaeus	1758	[46]
Larentiinae	Carsia sororiata	Hübner	1813	[23,34,36]
Larentiinae	Catarhoe cuculata	Hufnagel	1767	[37,46,59,63]
Larentiinae	Catarhoe rubidata	Denis & Schiffermüller	1775	[46]
Larentiinae	Chloroclysta miata	Linnaeus	1758	[36]
Larentiinae	Cidaria distinctata	Staudinger	1892	[37]
Larentiinae	Cidaria fulvata	Forster	1771	[34,44,63,65]
Larentiinae	Coenocalpe lapidata	Hübner	1809	[21,23,34,36,46,65]
Larentiinae	Coenotephria korschunovi	Viidalepp	1976	[34]
Larentiinae	Colostygia aptata	Hübner	1813	[34,65]
Larentiinae	Cosmorhoe ocellata	Linnaeus	1758	[37]
Larentiinae	Dysstroma citrata	Linnaeus	1761	[34,46,63,65]
Larentiinae	Dysstroma citrata septentrionalis	Heydemann	1929	[36]
Larentiinae	Dysstroma citratum kamtshadalarium	Beljaev & Vasilenko	2002	[48]
Larentiinae	Dysstroma infuscata	Tengström	1869	[65]
Larentiinae	Dysstroma latefasciata	Blöcker	1908	[34,44,65]
Larentiinae	Dysstroma pseudimmanata	Heydemann	1929	[31,34,44]
Larentiinae	Dysstroma truncata	Hufnagel	1767	[23,31,34,44,65,91]
Larentiinae	Dysstroma truncata transbaicalensis	Heydemann	1929	[36]
Larentiinae	Ecliptopera capitata	Herrich-Schäffer	1839	[63]
Larentiinae	Ecliptopera dimita	Prout	1938	[37]
Larentiinae	Ecliptopera umbrosaria	Motschulsky	1861	[34]
Larentiinae	Ecliptoptera oblongata	Guenée	1858	[44]
Larentiinae	Electrophaes chimakaleparia	Oberthür	1893	[44]
Larentiinae	Electrophaes corylata	Thunberg	1792	[46,65]
Larentiinae	Entephria caesiata	Denis & Schiffermüller	1775	[34,36,44]
Larentiinae	Entephria kuznetzovi	Viidalepp	1976	[34,45]
Larentiinae	Entephria tzygankovi	Wehrli	1929	[36]
Larentiinae	Epirrhoe alternata	Müller	1764	[23,34,36]
Larentiinae	Epirrhoe hastulata	Hübner	1790	[34,36,44,46]
Larentiinae	Epirrhoe hastulata reducta	Djakonov	1929	[48]
Larentiinae	Epirrhoe pupillata	Thunberg	1788	[23,34,36,44,46,63,65,91]
Larentiinae	Epirrhoe tristata	Linnaeus	1758	[23,34,46]
Larentiinae	Epirrita autumnata	Borkhausen	1794	[21,34,34]
Larentiinae	Epirrita autumnata smetanini	Beljaev & Vasilenko	2002	[48]
Larentiinae	Epirrita autumnata tunkunata	Bang-Haas	1910	[36]
Larentiinae	Esakiopteryx volitans	Butler	1878	[44]
Larentiinae	Eulithis mellinata	Fabricius	1787	[34]
Larentiinae	Eulithis populata	Linnaeus	1758	[36,44,63,91]
Larentiinae	Eulithis prunata	Linnaeus	1758	[34,44,46]
Larentiinae	Eulithis pyraliata	Denis & Schiffermüller	1775	[23,34,44,46,63,65]
Larentiinae	Eulithis pyropata	Hübner	1809	[91]
Larentiinae	Eulithis testata	Linnaeus	1761	[23,34,44,46,63]
Larentiinae	Euphyia coangulata	Prout	1914	[21,23,24,34,36,44,65]
Larentiinae	Euphyia intersecta	Staudinger	1882	[21,23,34]
Larentiinae	Euphyia unangulata	Haworth	1809	[34,46,63,65]
Larentiinae	Eupithecia selinata	Herrich-Schäffer	1861	[34]
Larentiinae	Eupithecia absinthiata	Clerck	1759	[95]
Larentiinae	Eupithecia actaeata	Walderdorff	1869	[52]
Larentiinae	Eupithecia addictata	Dietze	1908	[37]
Larentiinae	Eupithecia aggregata	Guenée	1858	[37]
Larentiinae	Eupithecia amplexata	Christoph	1881	[34,65]
Larentiinae	Eupithecia anikini	Mironov & Galsworthy	2014	[52]
Larentiinae	Eupithecia aporia	Vojnits	1975	[41,45]
Larentiinae	Eupithecia assimilata	Doubleday	1856	[52]
Larentiinae	Eupithecia bastelbergeri	Dietze	1910	[52]
Larentiinae	Eupithecia biornata	Christoph	1867	[34,65]
Larentiinae	Eupithecia bohatschi	Staudinger	1897	[25,34,65]
Larentiinae	Eupithecia carpophilata	Staudinger	1897	[34,65]
Larentiinae	Eupithecia catharinae	Vojnits	1969	[65]
Larentiinae	Eupithecia centaureata	Denis & Schiffermüller	1775	[34,63,65]
Larentiinae	Eupithecia chingana	Wehrli	1926	[45]
Larentiinae	Eupithecia corroborata	Dietze	1908	[36]
Larentiinae	Eupithecia denotata	Hübner	1813	[34]
Larentiinae	Eupithecia despectaria	Lederer	1853	[34,37]
Larentiinae	Eupithecia dissertata	Püngeler	1905	[34,36,65]
Larentiinae	Eupithecia djakonovi	Shchetkin	1956	[37]
Larentiinae	Eupithecia dolosa	Vojnits	1977	[45]
Larentiinae	Eupithecia ericeata	Rambur	1833	[52,65]
Larentiinae	Eupithecia extensaria	Freyer	1844	[36,65]
Larentiinae	Eupithecia fennoscandica	Knaben	1949	[36,96]
Larentiinae	Eupithecia fuscicostata	Christoph	1887	[65]
Larentiinae	Eupithecia graciliata	Dietze	1906	[34]
Larentiinae	Eupithecia hannemanni	Vojnits & De Laever	1973	[65]
Larentiinae	Eupithecia holti	Viidalepp	1973	[34,65,97]
Larentiinae	Eupithecia illaborata	Dietze	1904	[52]
Larentiinae	Eupithecia impolita	Vojnits	1980	[52]
Larentiinae	Eupithecia inculta	Vojnits	1975	[65]
Larentiinae	Eupithecia indigata	Hübner	1813	[63]
Larentiinae	Eupithecia innotata	Hufnagel	1767	[21,34,65]
Larentiinae	Eupithecia intricata	Zetterstedt	1839	[34]
Larentiinae	Eupithecia inveterata	Vojnits	1987	[65]
Larentiinae	Eupithecia irriguata	Hübner	1813	[65]
Larentiinae	Eupithecia kozlovi	Viidalepp	1973	[34,97]
Larentiinae	Eupithecia kuldschaensis	Staudinger	1892	[34,65]
Larentiinae	Eupithecia laboriosa	Vojnits	1977	[65]
Larentiinae	Eupithecia lariciata	Freyer	1841	[34,36,65]
Larentiinae	Eupithecia leptogrammata	Staudinger	1882	[65]
Larentiinae	Eupithecia linariata	Denis & Schiffermüller	1775	[65]
Larentiinae	Eupithecia mima	Mironov	1989	[65]
Larentiinae	Eupithecia minusculata	Alphéraky	1883	[34,65]
Larentiinae	Eupithecia mongolica	Vojnits	1974	[65]
Larentiinae	Eupithecia morosa	Vojnits	1976	[65]
Larentiinae	Eupithecia munguata	Mironov & Galsworthy	2014	[52]
Larentiinae	Eupithecia necessaria	Vojnits	1977	[41,45]
Larentiinae	Eupithecia nephelata	Staudinger	1897	[21,23,34,65]
Larentiinae	Eupithecia nobilitata	Staudinger	1882	[36,65]
Larentiinae	Eupithecia olgae	Mironov	1986	[52]
Larentiinae	Eupithecia opisthographata	Dietze	1906	[34]
Larentiinae	Eupithecia perfuscata	Vojnits	1975	[65]
Larentiinae	Eupithecia pernotata	Guenée	1858	[48]
Larentiinae	Eupithecia pimpinellata	Hübner	1813	[34,65]
Larentiinae	Eupithecia propria	Vojnits	1977	[65]
Larentiinae	Eupithecia pusillata	Denis & Schiffermüller	1775	[52]
Larentiinae	Eupithecia pygmaeata	Hübner	1799	[65]
Larentiinae	Eupithecia recens	Dietze	1904	[34,36]
Larentiinae	Eupithecia relaxata	Dietze	1904	[65]
Larentiinae	Eupithecia repentina	Vojnits & De Laever	1978	[52]
Larentiinae	Eupithecia rubellata	Dietze	1904	[41,45]
Larentiinae	Eupithecia saisanaria	Staudinger	1882	[52]
Larentiinae	Eupithecia satyrata	Hübner	1813	[36]
Larentiinae	Eupithecia selinata	Herrich-Schäffer	1861	[95]
Larentiinae	Eupithecia simpliciata	Haworth	1809	[52]
Larentiinae	Eupithecia sinuosaria	Eversmann	1848	[23,34,36]
Larentiinae	Eupithecia subbrunneata	Dietze	1904	[52]
Larentiinae	Eupithecia subexiguata	Vojnits	1974	[65]
Larentiinae	Eupithecia subfuscata	Haworth	1809	[34]
Larentiinae	Eupithecia suboxydata	Staudinger	1897	[65,98]
Larentiinae	Eupithecia subtacincta	Hampson	1895	[37]
Larentiinae	Eupithecia subumbrata	Denis & Schiffermüller	1775	[23,34,65]
Larentiinae	Eupithecia succenturiata	Linnaeus	1758	[95]
Larentiinae	Eupithecia sutiliata	Christoph	1877	[65]
Larentiinae	Eupithecia thalictrata	Püngeler	1902	[52]
Larentiinae	Eupithecia undata	Freyer	1840	[65]
Larentiinae	Eupithecia veratraria	Herrich-Schäffer	1848	[95]
Larentiinae	Eupithecia vicina	Mironov	1989	[65]
Larentiinae	Eupithecia virgaureata	Doubleday	1861	[21,23,34,65]
Larentiinae	Eupithecia vulgata	Haworth	1809	[21,23,34]
Larentiinae	Eupithecia vulgata lepsaria	Staudinger	1882	[37]
Larentiinae	Eupithecis unedonata	Mabille	1868	[33]
Larentiinae	Eustroma reticulatum obsoleta	Djakonov	1929	[48]
Larentiinae	Gagitodes sagittata	Fabricius	1787	[44,46,63]
Larentiinae	Gagitodes sagittata albiflua	Prout	1939	[48]
Larentiinae	Horisme aemulata	Hübner	1813	[23,34,46,65]
Larentiinae	Horisme aquata	Hübner	1813	[23,34,36,46,65,91]
Larentiinae	Horisme falcata	Bang-Haas	1907	[25,27,34,36,63,65]
Larentiinae	Horisme incurvaria	Erschoff	1877	[34,36,65]
Larentiinae	Horisme lucillata	Guenée	1858	[23,34]
Larentiinae	Horisme parcata	Püngeler	1909	[65]
Larentiinae	Horisme scotosiata	Guenée	1858	[21,23,34,63,65]
Larentiinae	Horisme tersata	Denis & Schiffermüller	1775	[34,65]
Larentiinae	Horisme tersata tetricata	Guenée	1858	[37]
Larentiinae	Horisme vitalbata	Denis & Schiffermüller	1775	[21,23,34,36,46,65]
Larentiinae	Hydrelia flammeolaria	Hufnagel	1767	[44,46]
Larentiinae	Hydria cervinalis	Scopoli	1763	[34]
Larentiinae	Hydria undulata	Linnaeus	1758	[34,65]
Larentiinae	Hydriomena furcata	Thunberg	1784	[21,23,34,36,44]
Larentiinae	Hydriomena impluviata	Denis & Schiffermüller	1775	[21,34,36]
Larentiinae	Hydriomena impluviata djakonovi	Beljaev & Vasilenko	2002	[48]
Larentiinae	Hydriomena ruberata	Freyer	1831	[65]
Larentiinae	Juxtephria consentaria	Freyer	1846	[36,44,65]
Larentiinae	Kyrtolitha obstinata	Staudinger	1892	[34]
Larentiinae	Laciniodes denigrata abiens	Prout	1938	[33]
Larentiinae	Lampropteryx albigirata	Kollar	1848	[65]
Larentiinae	Lampropteryx jameza	Butler	1898	[37]
Larentiinae	Lampropteryx minna	Butler	1881	[44,45,65]
Larentiinae	Lampropteryx suffumata	Denis & Schiffermüller	1775	[63]
Larentiinae	Leptostegna tenerata	Christoph	1881	[99]
Larentiinae	Lithostege coassata mongolica	Vojnits	1978	[42]
Larentiinae	Lithostege coassata ochraceata	Staudinger	1897	[42,65]
Larentiinae	Lithostege mesoleucata	Püngeler	1899	[34,42]
Larentiinae	Lithostege pallescens	Staudinger	1897	[21,34]
Larentiinae	Lobophora halterata	Hufnagel	1767	[44,46]
Larentiinae	Martania taeniata	Stephens	1831	[44]
Larentiinae	Mesoleuca albicillata	Linnaeus	1758	[34,37,44,46]
Larentiinae	Mesotype verberata	Scopoli	1763	[44]
Larentiinae	Nebula lamata	Staudinger	1897	[21,34]
Larentiinae	Nebula mongoliata	Staudinger	1897	[21,34,44,65]
Larentiinae	Odezia atrata	Linnaeus	1758	[23,34]
Larentiinae	Orthonama obstipata	Fabricius	1794	[34]
Larentiinae	Pelurga comitata	Linnaeus	1758	[34,44,63,65]
Larentiinae	Pelurga taczanowskiaria	Oberthür	1880	[63,91]
Larentiinae	Perizoma alchemillata	Linnaeus	1758	[34,36,44]
Larentiinae	Perizoma bifaciata	Haworth	1809	[65]
Larentiinae	Perizoma blandiata	Denis & Schiffermüller	1775	[23,34]
Larentiinae	Perizoma hydrata	Treitschke	1829	[36,44,65]
Larentiinae	Perizoma minorata	Treitschke	1828	[46]
Larentiinae	Phibalapteryx virgata	Hufnagel	1767	[34,36,42,91]
Larentiinae	Photoscotosia palaearctica	Staudinger	1882	[23,34]
Larentiinae	Plemyria rubiginata	Denis & Schiffermüller	1775	[34,44,65]
Larentiinae	Plesioscotosia pulchrata	Alphéraky	1883	[23,34]
Larentiinae	Povilasia kashghara	Moore	1878	[51]
Larentiinae	Pseudentephria remmi	Viidalepp	1976	[35]
Larentiinae	Pseudobaptria corydalaria	Graeser	1889	[34]
Larentiinae	Rheumaptera hastata	Linnaeus	1758	[34,36,44,46,65]
Larentiinae	Rheumaptera subhastata	Nolcken	1870	[36]
Larentiinae	Rheumaptera subhastata commixta	Matsumura	1925	[48]
Larentiinae	Schistostege nubilaria	Hübner	1799	[23,34,36,42,65]
Larentiinae	Scotopteryx chenopodiata	Linnaeus	1758	[23,34,46,63,65]
Larentiinae	Scotopteryx chenopodiata sibirica	Bang-Haas	1907	[42]
Larentiinae	Scotopteryx golovushkini	Kostjuk	1991	[65]
Larentiinae	Scotopteryx sinensis	Alphéraky	1883	[23,34]
Larentiinae	Scotopteryx transbaicalica	Djakonov	1955	[28,34,36]
Larentiinae	Spargania luctuata	Denis & Schiffermüller	1775	[23,34,44,63,65]
Larentiinae	Stamnodes danilovi	Erschoff	1877	[21,23,34,36,42,65]
Larentiinae	Stamnodes danilovi djakonovi	Alphéraky	1916	[33]
Larentiinae	Stamnodes pauperaria	Eversmann	1848	[65]
Larentiinae	Thera obeliscata	Hübner	1787	[34,91]
Larentiinae	Thera variata	Denis & Schiffermüller	1775	[23,34]
Larentiinae	Trichopterigia consobrinaria	Leech	1891	[44]
Larentiinae	Trichopteryx carpinata	Borkhausen	1794	[65]
Larentiinae	Xanthorhoe abrasaria	Herrich-Schäffer	1855	[36,44,65]
Larentiinae	Xanthorhoe deflorata	Erschoff	1877	[23,34,44,65]
Larentiinae	Xanthorhoe montanata	Denis & Schiffermüller	1775	[34,36,46]
Larentiinae	Xanthorhoe quadrifasiata tannuensis	Prout	1924	[45,63]
Larentiinae	Xanthorhoe sajanaria	Prout	1914	[36,44]
Larentiinae	Xanthorhoe sajanaria djakonovi	Vasilenko	1995	[100]
Larentiinae	Xanthorhoe spadicearia	Denis & Schiffermüller	1775	[44,46]
Larentiinae	Xanthorhoe stupida aridela	Prout	1937	[37]
Larentiinae	Zola terranea	Butler	1879	[34]
Sterrhinae	Cleta jacutica (Axel Hausmann: probably only one Cleta species occurring in Mongolia)	Viidalepp	1976	[36]
Sterrhinae	Cleta perpusillaria	Eversmann	1847	[65]
Sterrhinae	Cyclophora albipunctata	Hufnagel	1767	[46]
Sterrhinae	Cyclophora pendularia	Clerck	1759	[46]
Sterrhinae	Glossotrophia rufotinctata	Prout	1913	[49]
Sterrhinae	Holarctias rufinaria	Staudinger	1861	[58]
Sterrhinae	Idaea aureolaria	Denis & Schiffermüller	1775	[23,34,46]
Sterrhinae	Idaea biselata extincta	Staudinger	1897	[101]
Sterrhinae	Idaea muricata	Hufnagel	1967	[34]
Sterrhinae	Idaea muricata minor	Sterneck	1727	[40]
Sterrhinae	Idaea nitidata	Herrich-Schäffer	1861	[37]
Sterrhinae	Idaea nudaria	Christoph	1881	[37]
Sterrhinae	Idaea pallidata	Denis & Schiffermüller	1775	[34,40]
Sterrhinae	Idaea rufaria	Hübner	1799	[65]
Sterrhinae	Idaea rusticata	Denis & Schiffermüller	1775	[40,63]
Sterrhinae	Idaea serpentata	Hufnagel	1767	[23,34,36,41,63]
Sterrhinae	Idaea straminata	Borkhausen	1794	[34,91]
Sterrhinae	Idaea straminata sibirica	Djakonov	1926	[40]
Sterrhinae	Ochodontia adustaria	Fischer de Waldheim	1840	[23,34,65]
Sterrhinae	Rhodometra sacraria	Linnaeus	1767	[34]
Sterrhinae	Rhodostrophia jacularia	Hübner	1813	[21,23,34,36,40,41,63,65]
Sterrhinae	Rhodostrophia tyugui	Vasilenko	1998	[64]
Sterrhinae	Rhodostrophia ustyuzhanini	Vasilenko	2006	[49]
Sterrhinae	Rhodostrophia vibicaria	Clerck	1759	[34,46,63]
Sterrhinae	Scopula aequifasciata	Christoph	1881	[47]
Sterrhinae	Scopula albiceraria	Herrich-Schäffer	1847	[21,25,34,65]
Sterrhinae	Scopula albiceraria vitellinaria	Eversmann	1851	[40,41]
Sterrhinae	Scopula beckeraria	Lederer	1853	[34,40,41,63,65]
Sterrhinae	Scopula beckeraria amataria	Wehrli	1927	[36,40,65]
Sterrhinae	Scopula cajanderi	Herz	1903	[41,46]
Sterrhinae	Scopula caricaria	Reutti	1853	[46]
Sterrhinae	Scopula contramutata	Prout	1920	[34]
Sterrhinae	Scopula cumulata	Alpheraky	1883	[65]
Sterrhinae	Scopula decorata	Denis & Schiffermüller	1775	[21,23,34,41,63,65]
Sterrhinae	Scopula decorata przewalskii	Viidalepp	1975	[36,40,65]
Sterrhinae	Scopula dignata	Guenée	1858	[34]
Sterrhinae	Scopula floslactata	Haworth	1809	[37]
Sterrhinae	Scopula frigidaria	Möschler	1860	[47]
Sterrhinae	Scopula immorata	Linnaeus	1758	[23,34,36,40,46,63,65]
Sterrhinae	Scopula immutata contramutata	Prout	1913	[58]
Sterrhinae	Scopula impersonata	Walker	1861	[34]
Sterrhinae	Scopula impersonata macescens	Butler	1879	[40,41]
Sterrhinae	Scopula incanata	Linnaeus	1758	[34,41,65]
Sterrhinae	Scopula latelineata	Graeser	1892	[49]
Sterrhinae	Scopula marginepunctata	Goeze	1781	[23,34,63]
Sterrhinae	Scopula nigropunctata	Hufnagel	1767	[34]
Sterrhinae	Scopula nigropunctata subcandidata	Walker	1863	[37]
Sterrhinae	Scopula ornata	Scopoli	1763	[34,41,46]
Sterrhinae	Scopula permutata	Staudinger	1897	[34,39,65]
Sterrhinae	Scopula rubiginata	Hufnagel	1767	[34,40,41,63,65,91]
Sterrhinae	Scopula ternata	Schrank	1802	[25,34,36,46]
Sterrhinae	Scopula tessellaria	Boisduval	1840	[65]
Sterrhinae	Scopula umbelaria	Hübner	1813	[34,46,63]
Sterrhinae	Scopula umbelaria graeseri	Prout	1935	[41,65]
Sterrhinae	Scopula virgulata	Denis & Schiffermüller	1775	[23,34,40,41,46,63,65,91]
Sterrhinae	Scopula virgulata substrigaria	Staudinger	1900	[36]
Sterrhinae	Timandra griseata	Petersen	1902	[46]
Sterrhinae	Timandra paralias	Prout	1935	[34,40]
Sterrhinae	Timandra recompta	Prout	1930	[40,63]

). The most species-rich subfamily was Larentiinae with 203 species, while we recorded only one species in the subfamily Desmobathrinae. For 301 species with exact location data (Table S1), we recorded species richness within 2° × 2° grid cells in whole Mongolia (Figure 2).

Species richness was highest in the northern central part of the country, with 133 species recorded near Darkhan-Uul Aimag and the capital Ulaanbaatar. Four most frequently recorded species were Rhodostrophia jacularia (in n = 32 grids), Scopula beckeraria (n = 18) Scopula albiceraria (n = 17), and Horisme aquata (n = 17).

As a result of the Good–Turing theory, estimated species richness for whole Mongolia was 663.19 with 0.95 confidence interval (606.80–734.12), which is nearly double the observed species richness (Q2.est = 78.51; se = 32.31; Undetected # species= 275.19; Undetected percentage (%) = 41.49). Also, we constructed a sample-based interpolation and extrapolation curve of 301 species with exact reported location within 51 grids. The interpolated and extrapolated estimators of species richness show similar results (Figure 3), the curve was not asymptotic, indicating under-sampling of the communities.

In the next step we used the fourteen Mongolian ecoregions (Figure 1) to investigate the distribution of the sampled geometrid species in more detail. The most species-rich ecoregion was Daurian Forest Steppe with 142 species, while Khangai Mountains Alpine Meadow was the lowest in species richness with only three species of geometrid moths (Figure 4). One species (Rhodostrophia jacularia) occurred in 10 ecoregions, there were five further generalist species (Euphyia unangulata, Eupithecia nephelata, Scopula albiceraria, Scopula beckeraria) that occurred in eight to nine ecoregions. In contrast, 126 species were recorded only in one ecoregion. Four ecoregions were clearly under-sampled (Altai Alpine Meadow and Tundra, Dzungarian Basin Semi-Desert, Khangai Mountains Alpine Meadow, Sayan Alpine Meadows and Tundra) thus to avoid misleading interpretation, we excluded those ecoregions from further analysis.

Interpolation and extrapolation curves of particular ecoregions differ in their shapes, thus indicating different “sample quality”. Curves of Alashan Plateau Semi-Desert, Altai Montane Forest and Forest Steppe, Eastern Gobi Desert Steppe, Gobi Lakes Valley Desert Steppe, and Great Lakes Basin Desert Steppe are not asymptotic, only half of the estimated maximum species richness is sampled; while curves of Daurian Forest Steppe, Mongolian-Manchurian Grassland, Selenge-Orkhon Forest Steppe and Trans-Baikal Coniferous Forests are half asymptotic, thus tending to increase, while the curve of Sayan Montane Coniferous Forests is flattening, thus pointing to complete sampling of the moth community (Figure 5).

For assessment of beta-diversity, we calculated estimates of the abundance-based Sorensen Index between ecoregions (

Table 2. Pairwise estimates of similarity between ecoregions with online tool Spade [69]. Shown is the estimated abundance-based Sorensen Index. Colors refer to the map in Figure 1. Ecoregion abbreviations as in Figure 4. Highest and lowest values in bold.

C12(i,j)	Alashan	Altai	Daurian	Eastern	Gobi	Great	Mongolian	Sayan	Selenge	Trans
Alashan	1	0.504	0.184	0.595	0.716	0.446	0.433	0.097	0.206	0.244
Altai		1	0.451	0.64	0.742	0.702	0.523	0.311	0.594	0.445
Daurian			1	0.188	0.324	0.267	0.669	0.499	0.769	0.685
Eastern				1	0.942	0.644	0.533	0.127	0.424	0.141
Gobi					1	0.8	0.679	0.14	0.371	0.076
Great						1	0.497	0.301	0.544	0.139
Mongolian							1	0.417	0.719	0.522
Sayan								1	0.631	0.447
Selenge									1	0.606
Trans										1

). We excluded ecoregions with fewer than 20 species to avoid sampling bias in similarity analysis. The highest pairwise estimated Sorensen Similarity Index was between Eastern Gobi Desert Steppe and Gobi Lakes Valley Desert Steppe (β_s = 0.942), while the lowest were between Trans-Baikal Coniferous Forests and both of Gobi Lakes Valley Desert Steppe, Great Lakes Basin Desert Steppe (β_s = 0.076).

An NMDS ordination biplot (stress = 0.05) shows two separate groups of geometrid species communities within ecoregions (Figure 6). Altai Montane Forest and Forest Steppe, Alashan Plateau Semi-Desert, Eastern Gobi Desert Steppe, Gobi Lakes Valley Desert Steppe, and Great Lakes Basin Desert Steppe are clustered in the first group, Sayan Montane Coniferous Forests, Mongolian-Manchurian Grassland, Daurian Forest Steppe, Selenge-Orkhon Forest Steppe, and Trans-Baikal Coniferous Forests are grouped in the second group. Precipitation was positively correlated with NMDS1, while temperature was positively correlated with NMDS2, both correlations were highly significant (p < 0.01). Number of records was positively correlated with both axes but was not significant (

Table 3. NMDS vector fitted values. Temp: Max temperature of warmest month, Precipitation: Annual precipitation, Records: Number of records of geometrid moths.

Variable	NMDS1	NMDS2	r2	Pr (> 0)
Temperature	−0.32277	0.94648	0.7473	0.009
Precipitation	0.97252	−0.23281	0.9183	0.001
Records	0.73924	0.67344	0.5096	0.095

).

4. Discussion

In this study, we compiled a geometrid species checklist for Mongolia, examined species richness and diversity of geometrid communities among ecoregions. In addition, we investigated which environmental variables impact the distribution of geometrid moths. Compiling a species checklist on geometrid moths from a variety of sources published since 1892 was quite challenging, as names of species and locations were changing over the years, while sample efforts in different studies and areas differed considerably. Despite all our efforts we may not have included all species recorded in Mongolia in our list.

In total, we found 1973 records of 388 geometrid species of six subfamilies, but these records were not evenly sampled. The sample-based interpolation and extrapolation curve of gridded sample was not asymptotic, indicating that our records do not represent the whole potential geometrid fauna in Mongolia (Figure 3). Species richness for whole Mongolia was estimated as 663.19 species with Good–Turing theory and this estimated species richness was nearly double the observed species richness. These results confirm the rarefaction analysis and show that our inventory of geometrid moths in Mongolia is still incomplete, with less than 60% of the estimated species being recorded. The fact that countrywide diversity was highest in the grid cell of the capital draws further attention towards an obvious sampling bias with undersampling for the rest of the country. Moreover, we expect to find species of two other subfamilies, Orthostixinae and Alsophilinae in Mongolia. Species of these subfamilies were recorded in adjacent areas, such as in Kazakhstan and in China [37]. However, according to Müller et al. Alsophilinae is transferred to Ennominae, while the subfamily status of Orthostixinae is still not clear [62].

Given the huge size of Mongolia the estimated richness of 663 geometrid species for the whole country seems to be not high. But we wanted to compare the species richness of Mongolia with species richness of other countries similar in size. Norway + Sweden + Finland (1,173,940 km²) together are similar in size to Mongolia (1,564,000 km²). Altogether, for these countries, 341 geometrid species are recorded [84]. If we compare observed species richness (388) of Mongolia with the richness of those countries, it is almost similar; if we compare estimated species richness (663), it is almost double.

However, Scandinavia is an area at high latitudes, with harsh climate, not really suited for an ectotherm group like moths. Further south, Iberian Peninsula and Balearic Islands together, have 589 geometrid species (According to a personal information of Javier Gastón, one of the authors of the paper, due to scientific efforts the total number of Geometridae recorded on Iberian Peninsula and the Balearic Islands is now 605 species.) [85] and their areas (596,740 km² + 4564 km²) are almost three times smaller than the landlocked area of Mongolia, which is situated at higher latitude. Comparisons between distant countries are always somewhat lacking, but no figures on geometrid species richness are available for the countries in Inner Asia (e.g., Kazakhstan).

The most frequently recorded species, which occurred in 10 ecoregions of Mongolia, was Rhodostrophia jacularia, an inhabitant of steppe and semi-desert [34,86]. Sihvonen and Nupponen [87] studied female wing shape of this species, but we could not find other studies related to the biology of this species.

Most records were found in Daurian Forest Steppe, Selenge-Orkhon Forest Steppe, and Mongolian-Manchurian Grassland. For many ecoregions, rarefaction curves were not asymptotic, thus revealing that sampling there was incomplete. Two ecoregions have no geometrid moth records at all and were thus excluded from analysis, namely Khangai Mountains Conifer Forests and Sayan Intermontane Steppe. The less studied areas comprise higher altitude areas from central Mongolia, as well as border regions. Sampling in these ecoregions, many of them with high habitat heterogeneity, will certainly expand our checklist.

To assess beta diversity among these unevenly sampled groups we used an estimator for Soerenson similarity that includes unseen species in the calculation [70]. The results, on the one hand, reflect the high habitat heterogeneity of Mongolia, with is steep ecological north-south gradient and the diverse biomes of the country that promote high beta diversity (Table 1). On the other hand, it proved that ecoregions that include similar biomes had higher similarity of moth communities, a result corroborated by NMDS. The most similar ecoregions were Eastern Gobi Desert Steppe and Gobi Lakes Valley Desert Steppe that adjoin each other (β_s = 0.942).

In NMDS, ecoregions were grouped in two big groups. The first group included Alashan Plateau Semi-Desert, Eastern Gobi Desert Steppe, Gobi Lakes Valley Desert Steppe, Great Lakes Basin Desert Steppe and Altai Montane Forest and Forest Steppe, while in the second group there were Daurian Forest Steppe, Mongolian-Manchurian Grassland, Sayan Montane Coniferous Forests, Selenge-Orkhon Forest Steppe, and Trans-Baikal Coniferous Forests. The geographically nearest ecoregions were grouped together, and also the ecoregions included in the same group belonged to mostly same biome type (Table 1). The first group comprised mostly Deserts and Xeric Shrublands except Altai Montane Forest and Forest Steppe, while three ecoregions of the second group belonged to Temperate Grasslands, Savannas and Shrublands.

Environmental variables that shaped species distribution were nominated by forward selection in NMDS and included annual precipitation and maximum temperature of warmest quarter. Number of records was also selected as variable, but only temperature and precipitation were significant in NMDS, thus corroborating the general robustness of our analysis, which was less influenced by sample effort. The aforementioned groups of ecoregions in NMDS differ along the precipitation gradient and within groups in temperature, e.g., the montane forests regions of both groups have lower values of NMDS2.

In a study on Borneo, geometrid moths showed a similar relationship with precipitation and temperature [88]. Temperature has also been a major impact on geometrid species distribution in the Andes [89]. Moreover, habitat disturbance played a big role in shaping the geometrid moth ensemble in northern Borneo [90]. Similarly, grazing proved to be a factor influencing community pattern in Mongolian moths [4]. Temperature, rainfall and habitat disturbance are impacted by climate change and anthropgenic influence, so we expect future changes within the Mongolian geometrid communities. The species list we present here can be a tool helping to monitor these changes.

Finally, we have to admit that our study has a few weaknesses. We compiled records only from literature (we apologize if we missed any) due to limited time and funding. A total of 87 of the 388 species in our checklist are still missing an exact location. This information may be available in the museum collections pinned to the respective specimens. A detailed research in museums would have certainly brought more records and species. In addition, all our records were not systematically collected, which might affect the statistical analysis. The mere fact that data were sampled over a long period of time in different research projects, with different ways of sampling certainly impacts the value of a statistical analysis. For example, in our field study [4], we used UV light, but in other studies normal light bulbs were used, sometimes even moths have even been collected during day time. Together with the general problem of undersampling, these points hamper a more detailed analysis of the Mongolian geometrid communities at the present time.

Nevertheless, due to our study, future directions of research on Mongolian Geometridae have become more clear: geometrid moths are really under-studied in Mongolia. We found two unsampled and four extremely under-sampled ecoregions and for all ecoregions expected species numbers were higher than recorded ones. So, we expect to find many more amazing moth species in future collections in the respective regions.

5. Conclusions

In total, 1973 records of 388 species were recorded, but we also expect that many more species will be recorded in the future in more elaborated sampling designs, especially from locations of southern, eastern and western Mongolia. Despite the fact that our compiled data is not good enough to analyze the distribution and diversity pattern in full detail, our study could reveal the knowledge gaps and undersampled areas, provide a first estimate of the approximate species number in whole Mongolia (n = 663), visualize the currently recorded distribution and diversity pattern of geometrid moths of Mongolia and evaluate the main environmental factors that shape the communities.