ISSN 0974-7907 (Online) ISSN 0974-7893 (Print) Journal of Threatened Taxa

15 February 2019 (Online & Print) Vol. 11 | No. 2 | 13195–13250 PLATINUM 10.11609/jott.2019.11.2.13195-13250 OPEN www.threatenedtaxa.org ACCESS

Building evidence for conservation globally MONOGRAPH J TT ISSN 0974-7907 (Online); ISSN 0974-7893 (Print)

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continued on the back inside cover

Caption: front cover - funerarius © P.V. Cherenkov; back cover - © P.V. Cherenkov. Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250

Wasp fauna (Hymenoptera: , Chrysididae, , Monograph , , , Pompilidae, ,

Sphecidae, & Trigonalyidae) of Mordovia State ISSN 0974-7907 (Online) Nature Reserve and its surroundings in Russia ISSN 0974-7893 (Print) PLATINUM 1 2 Аlexander Ruchin & Аlexander Antropov OPEN ACCESS

1 Joint Directorate of the Mordovia State Nature Reserve and Smolny National Park,Dachnyi Lane, 4, Saransk, Republic of Mordovia 430011, Russia. 2 Zoological Museum of Moscow University, Bol’shaya Nikitskaya Ulitsa, 2, Moscow 125009, Russia. 1 [email protected] (corresponding author), 2 [email protected]

Abstract: Information is presented on 249 vespine species from 11 families in two regions of Mordovia (including Mordovia State Nature Reserve (MSNR)) and three districts of Nizhny Novgorod region (including Sarov) in Russia. Of these, 31 species are new to the Republic of Mordovia. Fauna of separate families of in Mordovia State Nature Reserve is represented by 220 species: one species in Bethylidae, 31 in Chrysididae, two in Tiphiidae, two in Mutillidae, one in Scoliidae, 28 in Pompilidae, 37 in Vespidae, nine in , 108 in Crabronidae, and one in Trigonalyidae. The biodiversity of wasps in the reserve is very high. Of the 283 species of wasps in Mordovia, 220 (77.7%) are found within the reserve. Species characteristics such as geographic distribution and certain aspects of biology are also provided. There is an obvious extension of the ranges of three species, Priocnemis fastigiata, deforme, and Scolia hirta, to the north. Keywords: Geographic distribution, Nizhny Novgorod region, range extension, Republic of Mordovia, Russia, Vespomorpha.

Russian abstract: Аннотация: В статье приведены сведения о 249 видах из 11 семейств осообразных двух районов Мордовии (включая Мордовский заповедник) и трех районов Нижегородской области (включая г. Sarov). Из них 31 вид является новым для Республики Мордовия. Фауна отдельных семейств ос Мордовского заповедника представлена 220 видами: Bethylidae – 1 вид, Chrysididae – 31, Tiphiidae – 2, Mutillidae – 2, Scoliidae – 1, Pompilidae – 28, Vespidae – 37, Sphecidae – 9, Crabronidae – 108, Trigonalyidae – 1. Биоразнообразие осообразных заповедника очень высокое. Из 283 видов ос Мордовии 220 видов (77,7%) обитают в его пределах. Приводятся такие характеристики видов, как географическое распространение, некоторые аспекты биологии. Отмечены факты расширения ареалов в северном направлении трех видов: Priocnemis fastigiata, Sceliphron deforme и Scolia hirta. Ключевые слова: Мордовский заповедник, Республика Мордовия, Нижегородская область, осы, фауна, Hymenoptera.

DOI: https://doi.org/10.11609/jott.4216.11.2.13195-13250 | ZooBank: urn:lsid:zoobank.org:pub:783951FE-11E9-453E-BCFE-7DF8CCC9E538

Editor: Arkady Lelej, Russian Academy of Sciences, Vladivostok, Russia. Date of publication: 15 February 2019 (online & print)

Manuscript details: #4216 | Received 22 April 2018 | Final received 08 February 2019 | Finally accepted 10 February 2019

Citation: Ruchin, A. & A. Antropov (2019). fauna (Hymenoptera: Bethylidae, Chrysididae, Dryinidae, Tiphiidae, Mutillidae, Scoliidae, Pompilidae, Vespidae, Sphecidae, Crabronidae & Trigonalyidae) of Mordovia State Nature Reserve and its surroundings in Russia. Journal of Threatened Taxa 11(2): 13195–13250; https:// doi.org/10.11609/jott.4216.11.2.13195-13250

Copyright: © Ruchin & Antropov 2019. Creative Commons Attribution 4.0 International License. JoTT allows unrestricted use, reproduction, and distribution of this article in any medium by adequate credit to the author(s) and the source of publication.

Funding: Joint Directorate of the Mordovia State Nature Reserve and Smolny National Park (Saransk, Russia); Zoological Museum of Moscow University (Moscow, Russia).

Competing interests: The authors declare no competing interests.

Author details: Аlexander B. Ruchin, Doctor of Sciences (in Biology), docent. Director of the Joint Directorate of the Mordovia State Nature Reserve and National Park “Smolny”. Honoured Ecologist of the Republic of Mordovia. Member of the Association Board of Directors of Russian Nature Reserves and National Parks. Member of the Public Chamber of the Republic of Mordovia. He consists in the Expert Council on Protected Areas of the Ministry of Natural Resources of Russia. He is a member of the Saint Petersburg Union of Scientists, the Society of Biotechnologists of Russia, the A.M. Nikolsky Herpetological Society, the Russian Entomological Society. Аlexander Antropov, Zoological Museum of Moscow University, кандидат биологических наук, старший научный сотрудник. He is a member of the Russian Entomological Society.

Author contribution:Both authors made equal contribution to a paper.

Acknowledgements: Many thanks to P.V. Cherenkov (Sarov), G.B. Semishin (Saransk), and L.V. Egorov (Cheboksary) for the help in collection of material and photographing, and to P. Rosa (Italy) for the help in identifying some Chrysididae species.

13195 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov

INTRODUCTION coniferous-broad leaved forest zone at the forest- steppe border. Forest communities occupy 89.3% of the Hymenoptera is one of the largest groups of , total territory. Vegetation cover of MSNR in general has including more than 153,000 species in the world fauna a taiga character with a definite gravitation towards a (Peters et al. 2017). This group includes Vespomorpha, nemoral complex with vegetation fluctuations. Forest- one of the largest infraorders that comprises several steppe elements are typical for this territory. Pine superfamilies and more than 92,000 species (Aguiar et Pinus sylvestris L. is the main forest-forming breed in al. 2013). At the same time, the distribution of a huge the reserve. It forms pure or mixed plant communities number of these species is still poorly understood. New in the southern, central, and western parts of MSNR. works are published all the time containing information Birch Betula pendula Roth and birch groves are the on range refinement of already-described species second largest forest communities in the reserve. These (Ponomarenko & Olmi 2007; Mokrousov 2011; Ćetković are predominantly secondary communities at the sites et al. 2011; Antropov & Mokrousov 2016; Nemkov 2016, of pine forest fellings and burnts. Young birch forests 2017a,b,c,d; Ruchin & Artaev 2016; Antropov 2017a,b; are especially abundant in the area that suffered from Danilov 2017; Danilov & Mokrousov 2017; Fateryga wildfire in 2010. Linden groves are located mainly in 2017; Lelej 2017; Lelej & Fadeev 2017; Loktionov & the northern part of the reserve. These are secondary Lelej 2017; Mokrousov 2017b; Mokrousov & Lelej plant communities that arose on the site of pine 2017; Rosa et al. 2017a) as well as descriptions of new forests and lime-spruce forests. Oak forests occupy a species from the European part of Russia (Loktionov relatively small area of the reserve. They are common et al. 2016; Mokrousov 2017a; Rosa et al. 2017b). On in the floodplain of Moksha in the western part of the the territory of the Republic of Mordovia (central part reserve. Spruce (formed mainly by Picea abies L.) and of Russia), however, this group of insects began to be alder forests (formed by Alnus glutinosa (L.) Gaertn.) studied actively only in recent years (Ruchin & Shibaev are located mainly in the floodplains of the rivers and 2008; Ruchin et al. 2009; Mokrousov et al. 2009, 2013; streams (Pushta, Vyaz-Pusta, Vorskliaj, and Arga) and Mokrousov 2010a,b; Ruchin & Kurmaeva 2010; Ruchin occupy small areas. Plant communities of small-leaved 2011, 2015; Ruchin & Lengesova 2012; Ruchin & Zryanin species (birch, aspen, and alder) are often formed on 2013; Ruchin & Antropov 2014, 2016; Ruchin & Egorov the sites of burnt forests (Khapugin et al. 2016; Ruchin & 2017). Makarkin 2017). The main areas of floodplain meadows The high species richness of this group and the mass are located along the river Moksha in the southwest of nature and diversity of its representative’s biology make MSNR . The territory of the reserve is nominally divided it one of the most significant among many terrestrial into quarters, which are numbered starting from the biocenoses. This article is devoted to the study of wasp north and then from west to east. There are forest fauna, a large group of stinging Hymenoptera, excluding houses (cordons) in some places of the reserve, often (: Apiformes) and (Formicoidea), situated next to large forest glades (Image 2). inhabiting Mordovia State Nature Reserve (MSNR) and The research material was collected from April its surroundings in Russia. to September in 2008, 2009, and 2011–2017 using the generally accepted entomologic methods of field research (Fasulati 1971). The bulk of the material was MATERIAL AND METHODS collected in the MSNR. Since the insects of the studied group are quite active and capable of moving, we also Mordovia State Nature Reserve is located in provide information about materials from the vicinity Temnikov District of the Republic of Mordovia on a of MSNR, namely, from the territories of Temnikov and forested right brink of the river Moksha and covers an Elniki districts of the Republic of Mordovia, as well as area of 321.62km2. From the north, the border runs Pervomaisk, Diveevsk, and Voznesenskoe districts of the along the river Satis — the right tributary of Moksha, Nizhny Novgorod region. All material was collected by further to the east — along the river Arga, which flows the first author (in other cases, the collector is indicated into the Satis. The western border runs along the rivers separately). The determination of the material was held Chernaya, Satis, and Moksha. From the south, the by A.V. Antropov. In total, more than 1,400 specimens forest-steppe approaches, naturally delineating the were studied. About 60 quarters of the reserve were boundary of the reserve (Image 1). investigated. The forestland of the reserve belongs to the mixed The following annotated list contains for each

13196 Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov

Figure 1. Map location of Mordovia State Nature Reserve and research sites (research sites of 2008, 2009, 2011–2017 are shown as red dots).

Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 13197 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov

A B

C D

E F

Image 1. Main biotopes of the Mordovia State Nature Reserve. A - floodplain meadow (quar. 401, © A.B. Ruchin), B - meadow near the Inorski cordon (© G.B. Semishin), C - sphagnum pine forest along sand dunes (quar. 336, © O.N. Artayev), D - birch forest (quar 420, © O.N. Artaev), E - glade (quar 446, © O.N. Artaev), F - pine forest with spruce (quar 383, © O.G. Grishutkin).

species the references to the literature on the species landscape, we didn’t indicate altitude above sea level from the territory of MSNR and its surroundings, when we indicated coordinates. New records for the the new collecting grounds in the reserve and the Republic of Mordovia are marked with an asterisk (*) surrounding area (unpublished data), date of collection, and new records for the territory of the reserve with two the number of specimens collected, the information on asterisks (**). The question mark symbol (?) indicates general distribution and biology (for some species), as the findings of species that require confirmation by well as remarks (if any). Due to the flat character of the modern material. The hyphen (-) symbol indicates the

13198 Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov species that were excluded from the wasp list fauna of Kushki (54029’49” N 43016’11”E). the reserve for various reasons that are indicated as General distribution: Russia (European part from remarks.The abbreviations used in the list are as follows: the west and the centre, Siberia, and Amur region), the RM - Republic of Mordovia, NNR - Nizhny Novgorod Mediterranean, Caucasus, Turkey, and Japan (Hokkaido, Region, MSNR - Mordovia State Nature Reserve, quar. Honshu, Shikoku, and Kyushu) (Rosa et al. 2017a). - quarter, cord. - cordon, Distr. - district, ex. - exemplar (specimen). Latin species names are provided according * Elampus constrictus (Förster, 1853) (= Elampus to online resources (Catalogue of Life website 2018; ambiguus Dahlbom, 1854) Chrysis.net 2018; Fauna Europaea 2018). Material: RM: 1 female, 18.vii.2016, MSNR, quar. 278 (54047’51”N & 43012’27”E); 1 female, 18.vii.2016, quar. 303 (54047’17”N & 43012’32”E); 1 female, 9.vi.2016, SPECIES LIST quar. 331 (54046’43”N & 43012’37”E). General distribution: Russia (the European part Superfamily from the northwest till Crimea and northern Caucasus, Family Bethylidae Urals, Altai, Buryatia, Transbaikalia, Amur region, and ? Epyris minor Kieffer, 1906 Primorsky Krai), , northern Africa, and northern Literature: Mokrousov et al. 2013. China (Móczár 1964; Rosa et al. 2017a). General distribution: Russia (the south of the Biology: It is a brood parasite of digger wasps European part), northern Africa, , Czech Mimesa (Crabronidae). Republic, , and (Macek et al. 2007; Lelej & Fadeev 2017). Elampus panzeri (Fabricius, 1804) Remark: The species requires additional refinement Material: NNR: 1 ex., 27.v.2016, Sarov (54055’08” N of range in the European part of Russia. 43017’40”E), coll. P.V. Cherenkov. General distribution: Russia (the whole European Family Chrysididae part, Siberia, and Primorsky Krai), Europe, Caucasus, Subfamily Cleptinae Turkey, and northeastern China (Rosa et al. 2017a). Cleptes nitidulus (Fabricius, 1793) Biology: It is a brood parasite of digger wasps Mimesa Literature: Mokrousov et al. 2013. bicolor (Jurine, 1807), M. equestris (Fabricius, 1804), and General distribution: Russia (the European part from M. lutaria (Fabricius, 1787) (Crabronidae). the northwest till Crimea and northern Caucasus, and western Siberia), Europe, and Turkey (Rosa et al. 2017a). Hedychridium coriaceum (Dahlbom, 1854) Biology: It is a parasite of the larvae of the Literature: Mokrousov et al. 2013; Ruchin & Nematus and Euura (). Antropov 2014. General distribution: Russia (the European part from Subfamily Chrysidinae the northwest till northern Caucasus and Urals), Europe, Tribe Elampini and Turkey (Rosa et al. 2017a). Elampus albipennis Mocsáry, 1889 Biology: It is a brood parasite of digger wasps Literature: Ruchin & Antropov 2016. Lindenius albilabris (Fabricius, 1793) (Crabronidae). Material: RM: 1 female, 18.vii.2016, MSNR, quar. 278 (54047’51”N & 43012’27”E); 1 female, 18.vii.2016, quar. Hedychridium elegantulum du Buysson, 1887 303 (54047’17”N & 43012’32”E); 1 female, 9.vi.2016, Literature: Mokrousov et al. 2013 (as Hedychridium quar. 331 (54046’43”N & 43012’37”E). elongatulum Du Buysson, 1887). General distribution: Russia (the European part from General distribution: Russia (the European part the centre and the south, including Crimea, Siberia, and from the centre till northern Caucasus) and European Primorsky Krai), eastern Europe, Saudi Arabia, United Mediterranean (Rosa et al. 2017a). Arab Emirates, Turkey, Turkmenistan, and Iran (Rosa et Biology: It is a probable brood parasite of digger al. 2017a). wasps Miscophus bicolor Jurine, 1807 (Crabronidae). Remark: The species is confirmed in MSNR. Hedychridium krajniki krajniki Balthasar, 1946 * Elampus bidens (Förster, 1853) Literature: Rosa et al. 2017b. Material: RM: 1 female, 2.vii.2016, Temnikov Distr., General distribution: Russia (the European part

Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 13199 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov from the centre till northern Caucasus) and European 1793) (Crabronidae). Mediterranean (Rosa et al. 2017a). * longicolle Abeille de Perrin, 1877 Hedychridium roseum (Rossi, 1790) Material: RM: 1 female, 19.vii.2015, Elniki Distr., Literature: Mokrousov et al. 2013. Novye Shaly (54042’13”N & 43038’25”E); NNR: 1 ex., General distribution: Russia (the whole European 18.vi.2015, Sarov (54055’08” N 43017’40”E), coll. P.V. part, Urals, Siberia, Amur region, Khabarovsk, and Cherenkov (Image 3). Primorye), Europe, Azerbaijan, Georgia, Abkhazia, and General distribution: Russia (the centre and the south northeastern China (Rosa et al. 2017a). of the European part, including Crimea and northern Biology: It is a brood parasite of digger wasps Astata Caucasus, Urals, western Siberia, and Primorsky Krai), boops (Schrank, 1781), Tachysphex pompiliformis west and south of Europe, north of Africa, Syria, Turkey, (Panzer, 1804), and, probably, Dryudella stigma Georgia, , Turkmenistan, and China (Rosa et (Panzer, 1809) and Harpactus tumidus (Panzer, 1801) al. 2017a). (Crabronidae). Biology: It is a brood parasite of digger wasps (Crabronidae). Hedychridium zelleri (Dahlbom, 1845) Literature: Mokrousov et al. 2013. Hedychrum nobile (Scopoli, 1763) General distribution: Russia (from the west and the Literature: Mokrousov et al. 2013; Ruchin & centre of the European part till northern Caucasus) and Antropov 2014. Europe (Rosa et al. 2017a). Material: RM: 2 female, 12.vii.2014, MSNR, quar. Biology: It is a brood parasite of digger wasps 86 (54053’38”N & 43035’59”E); 1 female, 1 male, Miscophus niger Dahlbom, 1844 and M. concolor 11.vii.2015, quar. 301 (54047’19”N & 43010’36”E); 1 Dahlbom, 1844 (Crabronidae). male, 16.vii.2015, quar. 442 (54043’56”N & 43015’02”E); 1 female, 18.vi.2016, quar. 115 (54051’46”N & * Hedychrum chalybaeum Dahlbom, 1854 43009’26”E); 1 female, 18.vii.2016, quar. 358 (54046’05”N Material: RM: 1 female, 18.vii.2016, MSNR, quar. 303 & 43011’44”E); 1 female, 20.vi.2015, Temnikov Distr., (54047’17”N & 43012’32”E); 1 female, 18.vii.2016, quar. Russkoe Karaevo (54040’55”N, 43013’52”E); 5 female, 9 331 (54046’43”N & 43012’37”E); 1 female, 23.vii.2016, male, 15.viii.2015, Purdoshki (54040’04”N & 43032’33”E); Elniki Distr., Novoyamskaya Sloboda (54044’24”N & 1 female, 1 male, 19.vii.2015, Elniki Distr., Novye Shaly 43042’59”E); 1 female, 23.vii.2016, Malye Mordovskie (54042’13”N & 43038’25”E). Poshaty (54041’00”N & 43043’50”E). General distribution: Russia (the European part General distribution: Russia (the European part, from the northwest till Crimea, Urals, Tuva, and Siberia), Siberia, Amur region, and Primorsky Krai), Europe, Europe, Caucasus, and Abkhazia (Rosa et al. 2017a). Mongolia, and central and northeastern China (Rosa et Biology: It is a brood parasite of digger wasps al. 2017a). arenaria (Linnaeus, 1758), C. rybyensis (Linnaeus, 1771), Biology: It is a brood parasite of digger wasps Cerceris C. quadrifasciata (Panzer, 1799), and C. quadricinta interrupta (Panzer, 1799) (Crabronidae). (Panzer, 1799) (Crabronidae) (Polidori et al. 2010).

Hedychrum gerstaeckeri Chevrier, 1869 Hedychrum rutilans Dahlbom, 1854 Literature: Mokrousov et al. 2013. Literature: Mokrousov et al. 2013. Material: RM: 1 female, 24.vii.2016, Temnikov Distr., Material: NNR: 1 ex., 23.vi.2016, Sarov (54055’08” N Polyanki (54044’27”N & 43029’04”E). 43017’40”E), coll. P.V. Cherenkov. General distribution: Russia (the European part, General distribution: Russia (the European part, Siberia, Amur region, Khabarovsk Krai, and Primorsky including Crimea and northern Caucasus, Urals, and Krai), Europe, Georgia, eastern China, and Japan (Rosa eastern Siberia), Europe, northern Africa, Turkey, et al. 2017a). Azerbaijan, and Georgia (Rosa et al. 2017a). Biology: It is a brood parasite of digger wasps Cerceris Biology: It is a brood parasite of digger wasps rybyensis (Linnaeus, 1771) and C. ruficornis (Fabricius, Philanthus triangulum (Fabricius, 1775) and P. coronatus

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(Thunberg, 1784) (Crabronidae) (Polidori et al. 2010). and the Middle East (Rosa et al. 2017a). Biology: It is a brood parasite of digger wasps Hedychrum virens Dahlbom, 1854 Passaloecus turionum Dahlbom, 1844, P. gracilis (Curtis, Literature: Ruchin & Antropov 2016. 1834), and P. eremita Kohl, 1893 (Crabronidae). Females Material: RM: 1 male, 12.vii.2014, Temnikov lay eggs on live aphids at the wasps’ hunting grounds, Distr., Alkaevo (54035’49”N & 43022’35”E); 1 female, but do not penetrate into their nests. 15.viii.2015, Purdoshki (54040’04”N & 43032’33”E). General distribution: Russia (the centre and the Philoctetes bidentulus (Lepeletier, 1806) south of the European part, including Crimea, Urals, and Literature: Mokrousov et al. 2013. western Siberia), west and south of Europe, the Middle General distribution: Russia (the European part, East, Turkey, Azerbaijan, and Georgia (Rosa et al. 2017a). including northern Caucasus, Urals, and Siberia), Biology: It is a brood parasite of digger wasps Cerceris European Mediterranean, northern Africa, the Middle tuberculata (de Villers, 1789) (Crabronidae). East, Turkey, and Caucasus (Rosa et al. 2017a). Biology: It is a brood parasite of digger wasps * Holopyga chrysonota (Förster, 1853) Psenulus pallipes (Panzer, 1798) (Crabronidae). Material: RM: 1 male, 16.vii.2016, MSNR, Pushta (54042’50”N & 43013’32”E). Philoctetes sareptanus (Mocsáry, 1889) General distribution: Russia (the European part, Literature: Ruchin & Antropov 2016. including Crimea and northern Caucasus, Urals, and Material: RM: 1 female, 27.v.2016, MSNR, quar. 360 Siberia), Mediterranean Europe, Caucasus, and northern (54046’13”N & 43013’31”E); 1 female, 11.vi.2016, quar. China (Rosa et al. 2017a). 364 (54046’24”N & 43017’45”E). Biology: It is a brood parasite of digger wasps General distribution: Russia (the European part, (Crabronidae). including northern Caucasus, Urals, Irkutsk region, and Khabarovsk Krai) and Iran (Rosa et al. 2017a). Omalus aeneus (Fabricius, 1787) Remark: The species is confirmed in MSNR. Literature: Mokrousov et al. 2009. Material: RM: 1 female, 13.vi.2016, MSNR, quar. 421 Philoctetes truncatus (Dahlbom, 1831) (54043’52’’N, 43007’06”E). Literature: Feoktistov 2011 (as Chrysellampus General distribution: Russia (the European part truncastus Dhlb.). from the northwest to the south, including Crimea General distribution: Russia (the centre and the east and northern Caucasus, Urals, Siberia, Amur region, of the European part, Urals, and Siberia), Europe, and Khabarovsk Krai and Primorsky Krai, Sakhalin, and Kuril northern Africa (Rosa et al. 2017a). Islands), Europe, northern Africa, Georgia, Uzbekistan, Biology: It is a brood parasite of digger wasps northern and eastern China, Japan (Hokkaido, Honshu, Diodontus tristis (Vander Linden, 1829) (Crabronidae). and Kyushu), and North America as a result of accidental importation (Rosa et al. 2017a). Pseudomalus auratus (Linnaeus, 1758) Biology: It is a brood parasite of digger wasps Literature: Ruchin & Antropov 2016. Passaloecus corniger Shuckard, 1837, P. eremita Kohl, General distribution: Russia (the European part from 1893, P. gracilis (Curtis, 1834), P. singularis Dahlbom, the northwest till Crimea and the northern Caucasus, 1844, P. turionum Dahlbom, 1844, Pemphredon lethifer Urals, Siberia, Kamchatka, and Primorsky Krai), Europe, (Shuckard, 1837), P. lugubris (Fabricius, 1793), and northern Africa, Caucasus, northern China, Korea, Japan Psenulus pallipes (Panzer, 1798) (Crabronidae). Females (Hokkaido, Honshu), and North America as a result of lay eggs on live aphids at the wasps’ hunting grounds, accidental delivery (Rosa et al. 2017a). but do not penetrate into their nests. Biology: It is a brood parasite of digger wasps Remark: The species is confirmed in MSNR. Passaloecus corniger Shuckard, 1837, P. eremita Kohl, 1893, P. insignis (Vander Linden, 1829), P. gracilis * Omalus biaccinctus (du Buysson, 1893) (Curtis, 1834), P. monilicornis Dahlbom, 1842, P. Material: RM: 1 female, 18.vii.2016, MSNR, quar. singularis Dahlbom, 1844, P. turionum Dahlbom, 1844, 331 (54046’43”N & 43012’37”E). Pemphredon inornata Say, 1824, P. lethifer (Shuckard, General distribution: Russia (the European part from 1837), P. lugens Dahlbom, 1842, P. lugubris (Fabricius, the northwest to the south, including Crimea), Europe, 1793), P. rugifer (Dahlbom, 1844), and Diodontus tristis

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(Vander Linden, 1829) (Crabronidae). Primorsky Krai, and Sakhalin), Europe, northern Africa, and Caucasus (Rosa et al. 2017a). Pseudomalus pussillus (Fabricius, 1804) Biology: It is a brood parasite of digger wasps Literature: Plavilshchikov 1964 (as Ellampus pusillus Tachysphex obscuripennis (Schenck, 1857), T. F.); Mokrousov et al. 2013. pompiliformis (Panzer, 1804), and Dinetus pictus Material: RM: 1 female, 5.vii.2016, MSNR, quar. 324 (Fabricius, 1793) (Crabronidae). (54046’28”N & 43005’40”E); 1 female, 21.v.2017, cord. Inorski (54044’15”N & 43008’53”E). * Chrysis cerastes Abeille, 1877 General distribution: Russia (the European part Material: RM: 1 female, 2.vii.2017, MSNR, cord. from the northwest till Crimea and northern Caucasus, Inorski (54044’15”N & 43008’53”E). Urals, and Siberia), Europe, northern Africa, Caucasus, General distribution: Russia (the European part and Kazakhstan (Rosa et al. 2017a). from the north till northern Caucasus and Altai), south Biology: It is a brood parasite of digger wasps of Europe, Azerbaijan, and Caucasus (Rosa et al. 2017a). Passaloecus eremita Kohl, 1893, P. insignis (Vander Biology: It is a brood parasite of solitary Vespidae Linden, 1829), and Pemphedron lethifer (Shuckard, Syneuodynerus egregius (Herrich-Schäffer, 1839) 1837) (Crabronidae). (Vespidae, Eumeninae). Remark: The species is confirmed in MSNR. ? Chrysis comparata Lepeletier de Saint Fargeau, 1806 Tribe Chrysidini Literature: Plavilshchikov 1964 (as Chrysis chevrieri ? Chrysidea pumila (Klug, 1845) Mock.). Literature: Plavilshchikov 1964 (as Chrysogona General distribution: Russia (the European part pumila Kl.). from the centre till Crimea and northern Caucasus, and General distribution: Russia (the European part Urals), south of Europe, northern Africa, and Caucasus from the centre to northern Caucasus and Urals), south (Rosa et al. 2017a). of Europe, and northern Africa (Rosa et al. 2017a). Remark: The species requires further research to be Remark: The species requires further research to be confirmed in Mordovia. confirmed in Mordovia. Chrysis fulgida Linnaeus, 1761 Chrysis analis Spinola, 1808 Literature: Ruchin & Antropov 2014, 2016. Literature: Mokrousov et al. 2009; Feoktistov 2011. General distribution: Russia (the European part General distribution: Russia (the European part from the northwest to northern Caucasus, Urals, Siberia, from the northwest till Crimea and northern Caucasus, Amur region, Kamchatka, and Primorsky Krai), Europe, and Urals), south of Europe, and Caucasus (Rosa et al. Caucasus, and northeastern China (Rosa et al. 2017a). 2017a). Biology: It is a brood parasite of the solitary Biology: It is a brood parasite of solitary wasps Osmia Vespidae allobrogus (de Saussure, 1955), andrenoides Spinola, 1808 (, Megachilinae). S. bifasciatus (Linnaeus, 1761), S. crassicornis (Panzer, 1798), S. murarius (Linnaeus, 1758) and, perhaps, Chrysis bicolor Lepeletier de Saint Fargeau, 1806 parietum (Linnaeus, 1758) (Vespidae, Literature: Mokrousov et al. 2013; Ruchin & Eumeninae). Antropov 2014. Material: RM: 1 female, 28.vii.2015, MSNR, quar. 79 Chrysis ignita (Linnaeus, 1758) (54053’11”N & 43029’18”E); 1 female, 12.vii.2014, quar. Literature: Redikortsev 1938; Plavilshchikov 1964; 86 (54053’38”N & 43035’59”E); 1 female, 11.vii.2015, Mokrousov et al. 2009, 2013; Ruchin & Antropov 2014. quar. 301 (54047’19”N & 43010’36”E); 1 female, Material: RM: 1 female, 31.v.2015, MSNR, quar. 345 16.viii.2015, quar. 324 (54046’28”N & 43005’40”E); 1 (54047’13”N & 43026’30”E); 1 female, 2.viii.2014, Pushta female, 4.vii.2015, quar. 368 (54046’37”N & 43021’45”E); (54042’50”N & 43013’32”E); 1 female, 12.vii.2017, cord. 1 female, 19.vii.2015, Elniki Distr., Novye Shaly Inorski (54044’15”N & 43008’53”E), coll. G.B. Semishin. (54042’13”N & 43038’25”E). General distribution: Russia (the European part General distribution: Russia (the European part from the northwest till Crimea and northern Caucasus, from the northwest till Crimea and northern Caucasus, Urals, Siberia, Amur region, Kamchatka, Khabarovsk and Urals, Siberia, Amur region, Khabarovsk Krai and Primorye, and Sakhalin), Europe, northern Africa, India,

13202 Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov northern China, and Korea (Rosa et al. 2017a). Eumeninae). Biology: It is a brood parasite of solitary Vespidae Ancistrocerus auctus (Fabricius, 1793), A. gazella Trichrysis cyanea (Linnaeus, 1758) (Panzer, 1798), A. nigricornis (Curtis, 1826),A . parietinus Literature: Mokrousov et al. 2009, 2013; Feoktistov (Linnaeus, 1761), A. parietum (Linnaeus, 1758), A. 2011; Ruchin & Antropov 2014. scoticus (Curtis, 1826), A. trifasciatus (Müller, 1776), Material: RM: 1 male, 13.vi.2016, quar. 422 and also, probably, Gymnomerus laevipes (Shuckard, (54043’55”N & 43007’39”E); NNR: 1 ex., 14.vi.2015, 1837), Euodynerus posticus (Herrich-Schäffer, 1841), Voznesenskoe Distr., Alamasovo (54054’03”N & Allodynerus delphinalis (Giraud, 1866), and Symmorphus 43008’56”E), coll. P.V. Cherenkov (Image 3). bifasciatus (Linnaeus, 1761) (Vespidae, Eumeninae). General distribution: Russia (the European part from the northwest till Crimea and northern Caucasus, Chrysis longula Abeille de Perrin, 1879 Urals, Siberia, Amur region, Kamchatka, Khabarovsk Krai Literature: Ruchin et al. 2009. and Primorsky Krai, and Sakhalin), Europe, northern General distribution: Russia (the European part Africa, central , northern China, Korea, and Japan from the northwest till Crimea and northern Caucasus, (Hokkaido & Honshu) (Rosa et al. 2017a). Urals, and Siberia), Europe, and central Asia (Rosa et al. Biology: It is a brood parasite of digger wasps 2017a). Trypoxylon figulus (Linnaeus, 1758) and Pemphredon Biology: It is a brood parasite of solitary Vespidae rugifer (Dahlbom, 1844) [Crabronidae], and Ancistrocerus antilope (Panzer, 1798), A. parietinus carbonarius (Scopoli, 1763) and Deuteragenia sp. (Linnaeus, 1761), Euodynerus posticus(Herrich-Schäffer, (Pompilidae). 1841), Symmorphus crassicornis (Panzer, 1798), and S. Remark: The species is confirmed in the territories murarius (Linnaeus, 1758) (Vespidae, Eumeninae). adjacent to MSNR.

* Chrysis rutilans Olivier, 1790 Tribe Parnopini Material: RM: 1 female, 22.vii.2017, Elniki Distr., Parnopes grandior (Pallas, 1771) Malye Mordovskie Poshaty (54041’00”N & 43043’50”E). Literature: Plavilshchikov 1964; Ruchin & Antropov General distribution: Russia (the European part from 2014, 2016; Ruchin & Egorov 2017; Ruchin et al. 2019. the northwest till Crimea and northern Caucasus, Urals, Material: RM: 1 female, 22.vii.2017, Elniki Distr., Siberia, and Primorsky Krai), Europe, northern Africa, Novye Shaly (54042’13”N & 43038’25”E). Caucasus, China, and Japan (Rosa et al. 2017a). General distribution: Russia (the European part Biology: It is a brood parasite of solitary Vespidae Gymnomerus laevipes (Shuckard, 1837), Jucancistrocerus from the northwest till Crimea and northern Caucasus, caspicus Giordani Soika, 1970, and Katamenes Urals, and Siberia), Europe, northern Africa, central Asia, flavigularis (Blüthgen, 1951) (Vespidae, Eumeninae). Azerbaijan, and Caucasus (Rosa et al. 2017a). Biology: It is a brood parasite of digger wasps Bembix * Pseudochrysis neglecta (Shuckard, 1836) rostrata (Linnaeus, 1758) (Crabronidae). Material: RM: 1 female, 31.v.2014, Elniki Distr., Malye Mordovskie Poshaty (54041’00”N & 43043’50”E). Family Dryinidae General distribution: Russia (the European part ? Gonatopus lunatus Klug, 1810 from the northwest till Crimea and northern Caucasus, Literature: Mokrousov et al. 2013. Urals, Siberia, and Khabarovsk Krai), Europe, Turkey, General distribution: Russia (the European part from , northwestern China, and North America the centre to northern Caucasus, Urals, and Siberia), (Rosa et al. 2017a). Europe, Morocco, the Middle East, Turkey, Kazakhstan, Biology: It is a brood parasite of solitary Vespidae central Asia, Mongolia, and Korea (Lelej & Loktionov Odynerus reniformis (Gmelin, 1790), O. spinipes 2017). (Linnaeus, 1758), and, probably accidentally, Biology: It is a parasitoid of [Cicadellidae, Ancistrocerus parietum (Linnaeus, 1758) and Deltocephalinae, and Aphrodinae] (Olmi 1994). Gymnomerus laevipes (Shuckard, 1837) (Vespidae, Remark: The species requires further research to be

Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 13203 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov confirmed in Mordovia. It is not yet registered in MSNR. adjacent to MSNR.

Superfamily Tiphia unicolor Lepeletier de Saint Fargeau, 1845 Family Tiphiidae Literature: Mokrousov et al. 2013. Tiphia femorata Fabricius, 1775 Material: RM: 1 female, 11.vii.2015, MSNR, quar. 301 0 0 Literature: Redikortsev 1938; Plavilshchikov 1964; (54 47’19”N & 43 10’36”E); 1 male, 21.vii.2013, quar. 0 0 Mokrousov et al. 2009, 2013; Feoktistov 2011; Ruchin 368 (54 46’37”N & 43 21’45”E); 1 female, 6.vii.2013, 0 0 & Antropov 2014. Temnikov Distr., Lavrentevo (54 29’44”N & 43 02’41”E); Material: RM: 1 female, 11.vii.2015, MSNR, quar. 1 female, 19.vii.2015, Elniki Distr., Malye Mordovskie 0 0 142 (54051’12”N & 43009’21”E); 1 female, 11.vii.2015, Poshaty (54 41’00”N & 43 43’50”E). General distribution: Russia (the centre and the quar. 301 (54047’19”N & 43010’36”E); 1 female, south of the European part, northern Caucasus, and 27.vii.2014, quar. 384 (54045’33”N & 43012’44”E); 3 Altai), European Mediterranean, and Turkey (Mokrousov female, 12.vii.2014, 24.viii.2017, quar. 86 (54053’38”N & & Lelej 2017). 43035’59”E); 1 female, 28.vii.2015, quar. 82 (54053’12”N Biology: It is a parasite of larvae of Scarabaeidae. & 43032’46”E); 5 female, 25.vii.2009, 23.viii.2017, 0 0 24.viii.2017, cord. Inorski (54 44’15”N & 43 08’53”E), Family Mutillidae coll. G.B. Semishin; 1 female, 19.vii.2016, quar. 330 Myrmosa atra Panzer, 1801 0 0 (54 46’44”N & 43 11’09”E); 1 female, 24.vii.2016, quar. Literature: Mokrousov et al. 2013. 0 0 399 (54 46’40”N & 43 27’27”E); 4 female, 24.viii.2017, Material: RM: 1 male, 4.vii.2015, MSNR, quar. 288 0 0 quar. 368 (54 46’37”N & 43 21’45”E); 2 female, 1 male, (54048’27”N & 43022’03”E); 2 male, 18.vii.2017, quar. 24.viii.2017, quar. 80 (54053’11”N & 43030’22”E); 1 female, 380 (54045’06”N & 43008’34”E). 5.ix.2017, Pushta (54042’50”N & 43013’32”E); 1 female, General distribution: Russia (the European part 12.vii.2014, Temnikov Distr., Alkaevo (54035’49”N & from the northwest till Crimea and northern Caucasus, 43022’35”E); 2 female, 24.vii.2016, Polyanki (54044’27”N Urals, and Siberia), Europe, Turkey, ? Iran, Kazakhstan, & 43029’04”E); 2 female, 19.vii.2015, 22.vii.2017, and Kyrgyzstan (Lelej 2017). Elniki Distr., Malye Mordovskie Poshaty (54041’00”N & Biology: It is a brood parasite of digger wasps 43043’50”E); 2 female, 19.vii.2015, 22.vii.2017, Novye Lindenius albilabris (Fabricius, 1793), L. panzeri (Vander Shaly (54042’13”N & 43038’25”E); 2 female, 23.vii.2016, Linden, 1829), Crabro peltarius (Schreber, 1784), Novoyamskaya Sloboda (54044’24”N & 43042’59”E); 2 palmipes (Linnaeus, 1767), C. wesmaeli female, 23.vii.2016, Cherlyaj (54045’53”N & 43040’58”E). (Vander Linden, 1829), Diodontus tristis (Vander General distribution: Russia (the European part, Linden, 1829), and Oxybelus uniglumis (Linnaeus, 1758) Urals, eastern Siberia, and Amur region), Europe, (Crabronidae). northern Africa, Turkey, Iran, Tajikistan, Kyrgyzstan, Kazakhstan, Mongolia, and North America as a result of Smicromyrme rufipes (Fabricius, 1787) Literature: Mokrousov et al. 2013. accidental importation (Mokrousov & Lelej 2017). Material: RM: 1 male, 3.vii.2016, MSNR, quar. 338 Biology: It occurs from July to September. Parasite (54047’06”N & 43019’40”E). of larvae of Amphimallon solstiale (Linnaeus, 1758), General distribution: Russia (the European part Anisoplia austriaca Herbst, 1783, and some other from the northwest till Crimea, Urals, and Siberia) (Lelej Scarabaeidae. 2017). Biology: It is a brood parasite of spider wasps Evagetes Tiphia minuta Vander Linden, 1827 [Pompilidae], of digger wasps Astata boops (Schrank, Literature: Ruchin & Antropov 2014. 1781), Tachysphex, Miscophus spurius (Dahlbom, Material: RM: 1 female, 23.vii.2016, Elniki Distr., 1832), Palarus variegatus (Fabricius, 1781), Oxybelus 0 0 Malye Mordovskie Poshaty (54 41’00”N & 43 43’50”E). bipunctatus Olivier, 1812, O. uniglumis (Linnaeus, General distribution: Russia (the European part 1758), Crossocerus wesmaeli (Vander Linden, 1829), from the northwest to northern Caucasus, Altai, Irkutsk and Cerceris arenaria (Linnaeus, 1758) (Crabronidae), as region, and Primorsky Krai) (Mokrousov & Lelej 2017). well as of solitary bees Halictus [Apidae, Halictinae] and Biology: It is a parasite of larvae of Scarabaeidae. Andrena (Apidae, Andreninae). Remark: The species is confirmed in the territories

13204 Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov

Family Scoliidae Antropov 2014. Scolia hirta Schrank, 1781 Material: RM: 1 female, 15.vi.2016, MSNR, quar. 429 Literature: Ruchin & Antropov 2016; Ruchin & (54044’31”N & 43014’59”E); 1 female, 28.vi.2015, quar. Artaev 2016. 249 (54049’39”N & 43032’08”E); 1 female, 16.vii.2017, Material: RM: 1 ex., 15.vii.2017, MSNR, Pushta cord. Podrubnyi (54047’51”N & 43008’48”E); 1 female, (54042’50”N & 43013’32”E); NNR: 6.viii.2014, 8.viii.2014, 19.v.2014, cord. Podrubnyi, 1 female, 6.viii.2017, cord. 9.ix.2017, Sarov (54055’08” N 43017’40”E), 4 ex., coll. Inorski (54044’15”N & 43008’53”E), coll. G.B. Semishin; P.V. Cherenkov (Image 3). 12 female, 19.v.2014, 18.vii.2014, 19.vii.2015, 5.vii.2016, General distribution: Russia (the European part from 21.vii.2016, 23.vii.2016, 5.viii.2016, 11.vii.2017, the centre till Crimea and northern Caucasus, Urals, 23.vii.2017, 24.vii.2017, 5.ix.2017, 18.ix.2017, Pushta and Siberia), Europe, Cyprus, Israel, Turkey, Caucasus, (54042’50”N & 43013’32”E). Abkhazia, Iran, Turkmenistan, Uzbekistan, Kazakhstan, General distribution: Russia (the European part, and Mongolia (Lelej & Mokrousov 2017; Mokrousov & Altai, western and eastern Siberia, Amur and Magadan Lelej 2017). regions, and Khabarovsk Krai and Primorsky Krai), Biology: The hosts are flower chafers Protaetia Europe, northwestern Africa, the Middle East, Turkey, morio (Fabricius, 1781), P. speciosissima (Scopoli, Iran, Kazakhstan, Tajikistan, Mongolia, northern China, 1786), P. splendidula (Faldermann, 1835), Cetonia Korea, and Japan (Hokkaido, Honshu, Shikoku, Kyushu, aurata (Linnaeus, 1761), Liocola lugubris Herbst, 1786, and Tsushima) (Loktionov & Lelej 2017). and Tropinota hirta (Poda, 1761) (Klausnitzer 2013). Biology: Nests are in the shape of separate oval Predators are parasitic of Conopidae. clay cells located in shelters under rocks, under bark, or in abandoned xylophagus tunnels. Prey: Family Pompilidae spiders from families Salticidae, Clubionidae, Subfamily Ceropalinae Oxyopidae, Gnaphosidae, , , Ceropales maculata (Fabricius, 1775) Lycosidae, Segestroodae, and . During Literature: Redikortsev 1938; Plavilshchikov 1964; transportation, the female usually separates the legs Mokrousov et al. 2013; Ruchin & Antropov 2016. of the victim. Predators: parasitoid wasps Picardiella Material: RM: 1 female, 18.vii.2016, MSNR, quar. 358 melanoleuca (Gravenhorst, 1829) (). (54046’05”N & 43011’44”E); 1 female, 24.vii.2016, quar. 398 (54046’22”N & 43025’48”E); 1 male, 19.vii.2016, Caliadurgus fasciatellus (Spinola, 1808) quar. 330 (54046’44”N & 43011’09”E); 1 female, Literature: Mokrousov et al. 2009, 2013. 6.viii.2017, quar. 419 (54045’39”N & 43022’50”E). Material: RM: 1 female, 17.viii.2014, MSNR, quar. General distribution: Russia (the European part 397 (54046’17”N & 43025’44”E); 1 female, 23.vii.2016, from the northwest till Crimea and northern Caucasus, Elniki Distr., Cherlyaj (54045’53”N & 43040’58”E); Urals, Siberia, Amur and Magadan regions, Chukotka, NNR: 1 female, 16.vii.2017, Voznesenskoe Distr., Khabarovsk Krai and Primorsky Krai, Sakhalin, and Kuril Zarya (54050’08”N & 43007’08”E); 1 male, 16.vii.2017, Islands), Europe, northern Africa, Cyprus, Armenia, Svobodnyi (54050’47”N & 43007’48”E). Azerbaijan, Georgia, Israel, Lebanon, Jordan, Turkey, General distribution: Russia (the European part Syria, Iran, Iraq, Turkmenistan, Kazakhstan, Kyrgyzstan, from the northwest to northern Caucasus, Urals, Altai, Mongolia, China, Korea, and Japan (Honshu) (Loktionov eastern Siberia, Amur region, Kamchatka, Khabarovsk & Lelej 2017). Krai and Primorsky Krai, and Sakhalin), Europe, Cyprus, Biology: Brood parasite of spider wasps Priocnemis Turkey, Kazakhstan, Kyrgyzstan, Mongolia, and North exaltata (Fabricius, 1775), Agenioideus cinctellus America as a result of introduction (Loktionov & Lelej (Spinola, 1807), infuscatus (Vander Linden, 2017). 1827), A. nigerrimus (Scopoli, 1763), A. viaticus (Linnaeus, Biology: Nests are in dense soil of open and forest 1758), anceps (Wesmael, 1851), A. rufa biotopes. Prey: spiders Meta (Tetragnathidae), Araneus (Haupt, 1927), Auplopus carbonarius (Scopoli, 1763), alsine (Walckenaer, 1802), A. diadematus (Linnaeus, and Episyron rufipes (Linnaeus, 1758) (Pompilidae). 1758), and Araniella cucurbitina (Clerck, 1757) (Araneidae). Subfamily Auplopus carbonarius (Scopoli, 1763) – Cryptocheilus octomaculatus (Rossi, 1790) Literature: Mokrousov et al. 2013; Ruchin & Literature: Plavilshchikov 1964 (as Pompilus

Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 13205 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov quadripunctatus F.). tunnels in wood. Prey: spiders Clubiona (Clubionidae). General distribution: Russia (the European part (the south, northern Caucasus, and Crimea)) (Tobias 1978); Priocnemis agilis (Shuckard, 1837) Europe (western, southern, and eastern), Morocco, Literature: Mokrousov et al. 2013. Algeria, Tunisia, Turkey, Cyprus, Syria, Israel, Iran, General distribution: Russia (the European part from Kyrgyzstan, Turkmenistan, and Tajikistan (Ghahari et al. the northwest till Crimea and northern Caucasus, Altai, 2014; Loktionov & Lelej 2017). and Irkutsk region), Europe, Cyprus, Turkey, Uzbekistan, Remark: It is distributed much more southerly. We Kazakhstan, and Kyrgyzstan. exclude this species from the fauna of the reserve and the region. Priocnemis cordivalvata Haupt, 1927 Literature: Redikortsev 1938 (Priocnemis Cryptocheilus versicolor (Scopoli, 1763) cordivalvatus Hpt.); Plavilshchikov 1964 (Priocnemis Literature: Mokrousov et al. 2013. cordivalvatus Hpt.); Mokrousov et al. 2013. Material: RM: 1 female, 15.viii.2015, Temnikov General distribution: Russia (the European part from Distr., Alkaevo (54035’49”N & 43022’35”E). the northwest to northern Caucasus, Krasnoyarsk Krai, General distribution: Russia (the European part from and Irkutsk region), Europe, central Asia, and Mongolia the centre till Crimea and northern Caucasus, and Urals), (Loktionov & Lelej 2017). Europe, Cyprus, Turkey, and Kyrgyzstan (Eversmann 1849; Tobias 1978; Loktionov & Lelej 2017). Priocnemis coriacea Dahlbom, 1843 Remark: The species is confirmed in the territories Literature: Ruchin et al. 2009. adjacent to MSNR. Material: RM: 1 male, 11.v.2016, MSNR, quar. 448 (54042’37”N & 43012’29”E). Deuteragenia bifasciata (Geoffroy, 1785) General distribution: Russia (the European part Literature: Ruchin & Antropov 2014 (as Dipogon from the centre till Crimea and northern Caucasus, and bifasciatus (Geoffroy, 1785)), 2016. Urals), Europe, and Turkey (Loktionov & Lelej 2017). Material: RM: 1 female, 14.vi.2016, MSNR, quar. 384 Biology: Prey: spiders Lycosidae and Gnaphosidae. (54045’33”N & 43012’44”E). Remark: The species is confirmed in MSNR. General distribution: Russia (the European part from the northwest till Crimea and northern Caucasus, Priocnemis exaltata (Fabricius, 1775) Urals, Siberia, Amur and Magadan regions, Khabarovsk Literature: Redikortsev 1938 (Priocnemis gibbus Krai and Primorsky Krai, and Sakhalin), Europe, Georgia, Scop.); Plavilshchikov 1964 (Priocnemis exultatus Pz., Kazakhstan, Kirghizia, Korea, and Japan (Hokkaido, Priocnemis gibbus Scop.); Mokrousov et al. 2013; Ruchin Honshu, Shikoku, and Kyushu) (Loktionov & Lelej 2017). & Antropov 2014. Biology: They build nests in rotten wood and Material: RM: 1 female, 18.vii.2016, MSNR, quar. abandoned burrows of other insects. Prey: spiders 331 (54046’43”N & 43012’37”E); 1 female, 9.viii.2014, Clubionidae, Dysteruidae, Salticidae, and Diaea dorsata quar. 413 (54045’12”N & 43016’54”E); 1 female, (Fabricius, 1777) (Thomisidae). 22.viii.2015, quar. 434 (54045’04”N & 43019’02”E); Remark: The species is confirmed in MSNR. 2 female, 11.vii.2015, cord. Podrubnyi (54047’51”N & 43008’48”E); 1 female, 13.viii.2016, quar. 446 Deuteragenia vechti Day, 1979 (54043’13”N & 43013’23”E); 1 female, 24.viii.2017, quar. Literature: Mokrousov et al. 2013; Ruchin & 368 (54046’37”N & 43021’45”E); 1 female, 24.vii.2016, Antropov 2016. Temnikov Distr., Polyanki (54044’27”N & 43029’04”E); Material: RM: 1 female, 10.vii.2014, MSNR, cord. 1 female, 23.vii.2016, Elniki Distr., Malye Mordovskie Inorski (54044’15”N & 43008’53”E), coll. G.B. Semishin. Poshaty (54041’00”N & 43043’50”E). General distribution: Russia (the European part General distribution: Russia (the European part from from the northwest to the southeast, Urals, Siberia, the northwest till Crimea, Urals, and Siberia). Europe, Amur and Magadan regions, Chukotka, Khabarovsk Krai Turkey, Iran, Kazakhstan, and Kyrgyzstan (Loktionov & and Primorsky Krai, Sakhalin, and Kuril Islands), Europe, Lelej 2017). Kazakhstan, Kyrgyzstan, and Japan (Hokkaido, Honshu) Biology: Nests in the soil, often in existing cavities. (Loktionov & Lelej 2017). Prey: spiders Lycosidae, Salticidae, and Pisauridae. Biology: They build nests in abandoned xylophagous

13206 Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov

* Priocnemis fastigiata Haupt, 1934 & 43001’19”E). Material: RM: 1 female, 6.viii.2017, MSNR, quar. General distribution: Russia (the European part 420 (54045’29”N & 43024’19”E); 1 female, 23.vii.2016, from the northwest till Crimea and northern Caucasus, Elniki Distr., Malye Mordovskie Poshaty (54041’00”N & Urals, Altai, and Irkutsk region), Europe, Cyprus, Turkey, 43043’50”E). Georgia, Iran, Uzbekistan, and Kyrgyzstan (Loktionov & General distribution: Russia (the centre and Lelej 2017). the southeast of the European part), European Biology: Nests in loose soil, often in existing Mediterranean, , Hungary, and Turkey (Straka cavities. Prey: spiders Trochosa terricola Thorell, 1856 2011; Loktionov & Lelej 2017). (Lycosidae), Gnaphosidae, and Thomisidae, which Remark: The range extends to the northwest. female wasps place into the prepared cavity in the soil (Edwards & Telfer 2002). Priocnemis gussakowskiji Wolf, 2004 Literature: Mokrousov et al. 2013 (as Priocnemis * Priocnemis pusilla Schiødte, 1837 confusor Wahis, 2006). Material: RM: 1 female, 7.vii.2016, MSNR, quar. 411 0 0 General distribution: Russia (the centre and, perhaps, (54 45’05”N & 43 14’54”E). the south of the European part), Europe, Jordan, Turkey, General distribution:Russia (the European part from and Kazakhstan (Loktionov & Lelej 2017). the northwest till Crimea and northern Caucasus, Omsk region, and Krasnoyarsk territory), Europe, Turkey, Priocnemis minuta (Vander Linden, 1827) Georgia, Iran, Kyrgyzstan, Turkmenistan, and Mongolia (Loktionov & Lelej 2017). Literature: Mokrousov et al. 2013. Biology: Nests in loose soil in existing cavities. Prey: General distribution: Russia (the European part from spiders Clubionidae and Salticidae. the centre till Crimea and northern Caucasus, Buryatia, Amur region, and Primorsky Krai), Europe, Turkey, Priocnemis schioedtei Haupt, 1927 Cyprus, Kazakhstan, Kyrgyzstan, and Turkmenistan Literature: Mokrousov et al. 2013. (Loktionov & Lelej 2017). General distribution: Russia (the European part from the northwest till Crimea and northern Caucasus), Priocnemis parvula Dahlbom, 1845 Europe, Turkey, Kazakhstan, and Mongolia (Loktionov & Literature: Mokrousov et al. 2013. Lelej 2017). General distribution: Russia (the European part Biology: Nests in loose soil in existing cavities. Prey: from the northwest till Crimea and northern Caucasus, spiders Clubionidae, Gnaphosidae, Salticidae, and Siberia, Amur and Magadan regions, Khabarovsk Krai Lycosidae (Edwards & Telfer 2002). and Primorsky Krai, and Sakhalin), Europe, Tunisia, Jordan, Turkey, Uzbekistan, and Kyrgyzstan (Loktionov * Priocnemis vulgaris (Dufour, 1841) & Lelej 2017). Material: RM: 1 female, 11.vi.2016, MSNR, quar. 389 Biology: Nests in sandy soil. Prey: spiders Pardosa (54045’50”N & 43017’50”E); 4 female, 13.vi.2016, quar. prativaga (L.Koch, 1870), P. pullata (Clerck, 1757), 435 (54043’41”N & 43007’59”E); 1 female, 26.vi.2016, Xerolycosa miniata (C.L. Koch, 1834) [Lycosidae], quar. 403 (54044’40”N & 43006’57”E); 1 female, Haplodrassus soerenseni (Strand, 1900) (Gnaphosidae), 13.vi.2016, Pushta (54042’50”N & 43013’32”E). and Evarcha arcuata (Clerck, 1757) (Salticidae). General distribution: Russia (the European part from the northwest to northern Caucasus), south of Europe, Priocnemis perturbator (Harris, 1780) Turkey, Iran, Kazakhstan, Uzbekistan, Kyrgyzstan, and Literature: Ruchin et al. 2009; Mokrousov et al. Tajikistan (Loktionov & Lelej 2017). 2013; Ruchin & Antropov 2014. Material: RM: 1 female, 25.v.2015, MSNR, quar. 342 Subfamily (54046’21”N & 43023’23”E); 1 female, 10.v.2015, quar. (Dahlbom, 1829) 368 (54046’37”N & 43021’45”E); 3 female, 18.v.2014, Literature: Mokrousov et al. 2013. 7.vi.2015, quar. 399 (54046’40”N & 43027’27”E); 1 General distribution: Russia (the European part female, 6.v.2015, quar. 431 (54044’31”N & 43017’14”E); 1 from the northwest till northern Caucasus, Altai, eastern female, 16.v.2015, quar. 408 (54044’57”N & 43011’47”E); Siberia, Amur region, and Khabarovsk Krai and Primorsky 1 female, 6.vi.2015, Temnikov Distr., Vesely (54033’09”N Krai), Europe, Cyprus, Israel, Turkey, Iran, Kazakhstan,

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Uzbekistan, Kyrgyzstan, Turkmenistan, and Mongolia Plavilshchikov 1964 (as Anoplius fuscus F.); Mokrousov (Loktionov & Lelej 2017). et al. 2009; Feoktistov 2011; Ruchin & Antropov 2014. Biology: Nests in sandy soil, often under rocks. Material: RM: 1 female, 20.iv.2014, MSNR, quar. Prey: spiders Arctosa stigmosa (Thorell, 1875), Pardosa 434 (54045’04”N & 43019’02”E); 1 female, 27.v.2016, amentata (Clerck, 1757), P. lugubris (Walkenaer), Pirata quar. 360 (54046’13”N & 43013’31”E); 1 female, hygrophilus Thorell, 1872, P. piraticus (Clerck, 1757), 15.v.2017, quar. 302 (54047’15”N & 43011’26”E); 2 Trochosa ruricola (De Geer, 1778), T. terricola Thorell, male, 1.vii.2016, quar. 379 (54045’17”N & 43007’51”E); 1856 (Lycosidae), Dolomedes fimbriatus (Clerck, 1757) 1 female, 13.viii.2016, quar. 446 (54043’13”N & (Pisauridae), Heliophanus auratus C.L.Koch, 1835, 43013’23”E); 3 female, 11.v.2015, 14.V.2017, cord. H. dubius C.L. Koch, 1835, and Sitticus distinguendus Inorski (54044’15”N & 43008’53”E); 1 female, 12.v.2013, (Simon, 1868) (Salticidae). Temnikov Distr., Kitsaevka (54036’04”N & 43010’03”E); 1 male, 24.vii.2016, Polyanki (54044’27”N & 43029’04”E); 1 (Vander Linden, 1827) male, 22.vii.2017, Elniki Distr., Novye Shaly (54042’13”N Literature: Mokrousov et al. 2013. & 43038’25”E). Material: RM: 1 female, 23.viii.2017, MSNR, cord. General distribution: Russia (the European part Inorski (54044’15”N & 43008’53”E). from the northwest till Crimea and northern Caucasus, General distribution: Russia (the European part Urals, Siberia, Amur region, Khabarovsk and Primorye from the northwest till Crimea and northern Caucasus, territories, and Sakhalin), Europe, Cyprus, Egypt, Urals, Irkutsk Region, Amur region, Khabarovsk Krai Jordan, Turkey, Georgia, Iran, Kazakhstan, Uzbekistan, and Primorsky Krai, Sakhalin, and Kuril Islands), Europe, Kyrgyzstan, Turkmenistan, Mongolia, northern China, Cyprus, Algeria, Egypt, Jordan, Iran, Turkmenistan, ? Korea, and Japan (Hokkaido, Honshu, and Kyushu) Uzbekistan, Kyrgyzstan, Mongolia, northern China, (Loktionov & Lelej 2017). Korea, and Japan (Hokkaido, Honshu, Shikoku, Kyushu, Biology: Prey: spiders Alopecosa pulverulenta and Ryukyu) (Loktionov & Lelej 2017). (Clerck, 1775), A. (Clerck, 1775), A. cuneata Biology: Nests in moist sandy soil. Prey: spiders (Clerck, 1775), Dolomedes, (Clerck, Trochosa ruricola (De Geer, 1778) (Lycosidae) and 1775) (Pisauridae), Arctosa, Pardosa paludicola (Clerck, (Thomisidae). 1775), P. lugubris (Walckenaer, 1802), P. palustris Remark: The species is confirmed in MSNR. (Linnaeus, 1758), Pirata piraticus (Clerck, 1757), Trochosa terricola Thorell, 1856, T. ruricola (De Geer, (Scopoli, 1863) 1778), T. spinipalpis (F.O. Pickard-Cambridge, 1895) Literature: Redikortsev 1938 (as Apoplous nigerrimus (Lycosidae), Drassodes lapidosus (Walckenaer, 1802), Scop.); Plavilshchikov 1964; Mokrousov et al. 2013. Haplodrassus signifer (C.L. Koch, 1839) (Gnaphosidae), Material: RM: 1 female, 24.vii.2016, MSNR, quar. Aelurillus, Evarcha arcuata (Clerck, 1757) (Salticidae), 398 (54046’22”N & 43025’48”E). Agelena (Agelenidae), Agroeca brunnea (Blackwall, General distribution: Russia (the European part 1833) (Liocranidae), and Thanatus formicinus (Clerck, from the northwest till Crimea and northern Caucasus, 1757) (Philodromidae). Urals, Altai, eastern Siberia, Amur region, Kamchatka, Khabarovsk Krai and Primorsky Krai, Sakhalin, and Kuril (Wesmael, 1851) Islands), Europe, Turkey, Iran, Uzbekistan, Tajikistan, Literature: Mokrousov et al. 2013. and North America as a result of accidental delivery General distribution: Russia (the European part from (Loktionov & Lelej 2017). the northwest till Crimea and northern Caucasus, Altai, Biology: Nests in the soil in existing cavities (hollow eastern Siberia, Amur and Magadan regions, Kamchatka, stems of plants, abandoned nests of insects, cracks in the and Chukotka), Europe, Turkey, Syria, Iran, Kazakhstan, soil, and empty shells of snails) or simply in the thickness Kyrgyzstan, and Mongolia (Loktionov & Lelej 2017). of moss cover. Prey: spiders Pardosa, Trochosa ruricola Biology: Nests in loose soil. Prey: spiders Agelenidae, (De Geer, 1778) (Lycosidae), Clubionidae, Gnaphosidae, Clubionidae, Gnaphosidae, Lycosidae, Salticidae, and and Pisauridae. Thomisidae. Predators: brood parasites Ceropales Remark: The species is confirmed in MSNR. maculata (Fabricius, 1775) and (Shuckard, 1837). Anoplius viaticus (Linnaeus, 1758) Literature: Redikortsev 1938 (as Apoplous fuscus F.);

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* Arachnospila minutula (Dahlbom, 1842) Mongolia. Material: RM: 1 female, 13.vi.2016, MSNR, Pushta Biology: Prey: spiders Agelenidae, Clubionidae, (54042’50”N & 43013’32”E). Gnaphosidae, Lycosidae, Salticidae, and Thomisidae. General distribution: Russia (the European part Remark: The species requires further research to be from the northwest till Crimea and northern Caucasus, confirmed in Mordovia. Urals, Altai, Buryatia, Transbaikalia, Amur region, and Khabarovsk Krai and Primorsky Krai), Europe, the Middle Parabatozonus lacerticida (Pallas, 1771) East, Turkey, Kazakhstan, Kyrgyzstan, and Mongolia Literature: Redikortsev 1938 (as Batozonus (Loktionov & Lelej 2017). lacerticida Pall.); Plavilshchikov 1964 (as Batozonus Biology: Nests in loose soil. Prey: Agelenidae, lacerticida Pall.); Ruchin & Antropov 2014 (as Clubionidae, Gnaphosidae, Lycosidae, Salticidae, and (Pallas, 1771)). Thomisidae. Material: RM: 1 female, 11.vii.2015, MSNR, quar. 116 (54052’02”N & 43010’25”E); 1 female, 28.vi.2015, cord. Arachnospila spissa (Schioedte, 1837) Inorski (54044’15”N & 43008’53”E); 1 female, 23.vii.2016, Literature: Mokrousov et al. 2013. Elniki Distr., Malye Mordovskie Poshaty (54041’00”N Material: RM: 1 female, 16.vii.2015, MSNR, quar. & 43043’50”E); NNR: 2 ex., 27.vii.2014, Voznesenskoe 442 (54043’56”N & 43015’02”E); 1 female, 15.vii.2017, Distr., Alamasovo (54054’03”N & 43008’56”E), coll. P.V. quar. 324 (54046’28”N & 43005’40”E). Cherenkov. General distribution: Russia (the European part General distribution: Russia (the European part from from the northwest to northern Caucasus, Altai, eastern the northwest till Crimea and northern Caucasus, Urals, Siberia, Amur and Magadan regions, and Sakhalin), Altai, eastern Siberia, Amur region, and Khabarovsk Krai Europe, the Middle East, Turkey, Kazakhstan, central and Primorsky Krai), Europe, Morocco, Cyprus, Turkey, Asia, Mongolia, and northern China (Loktionov & Lelej Georgia, Syria, Iran, Kazakhstan, Uzbekistan, Kirghizia, 2017). Turkmenistan, Mongolia, northeastern China, Korea, Biology: Prey: spiders Lycosidae and Salticidae. and Japan (Hokkaido and Honshu) (Loktionov & Lelej Paralyzed prey is left in the burrow of a spider or hidden 2017). in the crevices of tree trunks or stumps. Biology: Nests in sandy soil. Prey: spiders Araneus and Argiope (Araneidae). * Arachnospila trivialis (Dahlbom, 1843) Material: RM: 1 female, 4.vii.2017, MSNR, quar. 357 Episyron rufipes (Linnaeus, 1758) (54046’07”N & 43010’56”E); 1 female, 9.vi.2016, quar. Literature: Mokrousov et al. 2013; Ruchin & 331 (54046’43”N & 43012’37”E); 1 female, 23.vii.2016, Antropov 2016. Elniki Distr., Malye Mordovskie Poshaty (54041’00”N & General distribution: Russia (the European part 43043’50”E). from the northwest till Crimea and northern Caucasus, General distribution: Russia (the European part from Urals, eastern Siberia, Amur and Magadan regions, the northwest to northern Caucasus, Altai, Amur and Kamchatka, Khabarovsk Krai and Primorsky Krai, and Magadan regions, Kamchatka, and Chukotka), Europe, Sakhalin), Europe, Morocco, Turkey, Cyprus, Sudan, the Middle East, Turkey, Kazakhstan, Kyrgyzstan, Ethiopia, Jordan, Yemen, Saudi Arabia, Syria, Iran, Tajikistan, and Mongolia (Loktionov & Lelej 2017). Kazakhstan, Uzbekistan, Kyrgyzstan, and Turkmenistan Biology: Nests in the soil. Prey: spiders Agelenidae, (Loktionov & Lelej 2017). Clubionidae, Gnaphosidae, Lycosidae, Salticidae, and Biology: Nests in sandy soil. Prey: spiders, mainly Thomisidae. Predators: brood parasites Evagetes Meta and Araneus [Araneidae], and sometimes crassicornis (Shuckard, 1837) (Pompilidae). Lycosidae.

? Arachnospila wesmaeli (Thomson, 1870) Evagetes crassicornis (Shuckard 1837) Literature: Redikortsev 1938 (as Psammochares Literature: Mokrousov et al. 2013. wesmaeli Thoms.); Plavilshchikov 1964 (as General distribution: Russia (the European part Psammochares wesmaeli Thms.). from the northwest to the centre and the east, Altai, General distribution: Russia (the European part eastern Siberia, Amur and Magadan regions, Chukotka, from the northwest to the centre and, probably, to the and Sakhalin), Europe, Morocco, Turkey, Georgia, south, and Krasnoyarsk Krai), Europe, Kyrgyzstan, and Kazakhstan, Kyrgyzstan, Mongolia, and North America

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(Loktionov & Lelej 2017). of plants, on clay slopes, and in old nests of other Biology: Brood parasite of spider wasps Arachnospila Hymenoptera). Prey: caterpillars of various Lepidoptera, anceps (Wesmael, 1851), A. consobrina (Dahlbom, 1843), less often false larvae of sawflies (Tenthredinidae), A. minutula (Dahlbom, 1842), A. trivialis (Dahlbom, larvae of snout beetles (Curculionidae), and leaf beetles 1843), and Anoplius nigerrimus (Scopoli, 1863) (Edwards (Chrysomelidae). Predators: emerald wasps Chrysis & Telfer 2002). longula Abeille de Perrin, 1879 and Ch. pseudobrevitarsis Linsenmaier, 1951 (Chrysididae). Homonotus sanguinolentus (Fabricius, 1793) Literature: Mokrousov et al. 2013. Ancistrocerus auctus (Fabricius 1793) General distribution: Russia (the European part Literature: Mokrousov et al. 2013. from the northwest till Crimea and Irkutsk and Magadan General distribution: Russia (the European part regions), Europe, Cyprus, Turkey, Syria, Iran, Kazakhstan, from the centre till Crimea and northern Caucasus and Kirghizia, and Turkmenistan (Loktionov & Lelej 2017). Urals), south of Europe, Israel, Jordan, Turkey, Caucasus, Biology: Prey: spiders Chiracanthium (Clubionidae). Kazakhstan, central Asia, Syria, Iraq, and Iran (Fateryga 2017). * Pompilus cinereus (Fabricius, 1775) Biology: Nests in existing cavities (in the stems Material: RM: 1 female, 2.vii.2016, Temnikov Distr., of plants, on clay slopes, and in old nests of other Kushki (54029’49” N 43016’11”E). Hymenoptera). Prey: caterpillars of various Lepidoptera, General distribution: Russia (the European part less often false larvae of sawflies (Tenthredinidae) and from the northwest till Crimea and northern Caucasus, larvae of snout beetles (Curculionidae) and leaf beetles Amur region, and Primorsky Krai), Europe, Algeria, (Chrysomelidae). Predators: emerald wasps Chrysis Tunisia, Egypt, Israel, Cyprus, Turkey, , northern ignita (Linnaeus, 1758) (Chrysididae). and southeastern China, Korea, Japan (Honshu, Kyushu, and Ryukyu), and Afrotropical, Oriental, and Australian Ancistrocerus gazella (Panzer, 1798) regions (Loktionov & Lelej 2017). Literature: Redikortsev 1938 (as Odynerus gazella Biology: Nests in sandy soil. Prey: spiders Panz.); Plavilshchikov 1964 (as Odynerus gazella Panz.). Zoropsis [Zoropsidae], Aelurillus [Salticidae], Xysticus Material: RM: 1 male, 16.vii.2015, MSNR, quar. 442 [Thomisidae], Chiracanthium, Zora [Clubionidae], (54043’56”N & 43015’02”E). Arctosa, Alopecosa, Lycosa, Xerolycosa, Pirata, Trochosa General distribution: Russia (the European part from [Lycosidae], Araneus, and Dolomedes (Araneidae). the centre till Crimea and northern Caucasus), Europe, northern Africa, Israel, Cyprus, Turkey, Caucasus, Iran, Family Vespidae Afghanistan, and ? Pakistan. Acclimatized in North Subfamily Eumeninae America and New Zealand as a result of accidental Ancistrocerus antilope (Panzer, 1798) importation (Fateryga 2017). Literature: Redikortsev 1938 (as Odynerus antilope Biology: Nests in existing cavities (in the stems Panz.); Plavilshchikov 1964 (as Odynerus antilope Panz.); of plants, on clay slopes, and in old nests of other Mokrousov et al. 2013; Ruchin & Antropov 2014. Hymenoptera). Prey: caterpillars of various Lepidoptera, Material: RM: 1 female, 13.v.2014, MSNR, quar. 360 less often false larvae of sawflies Tenthredinidae and (54046’13”N & 43013’31”E); 1 female, 9.vi.2016, quar. larvae of Curculionidae and Chrysomelidae. Predators: 331 (54046’43”N & 43012’37”E); 1 male, 31.v.2014, emerald wasps Chrysis ignita (Linnaeus, 1758) and Ch. cord. Taratinski (54044’43”N & 43005’14”E); 1 female, schencki Linsenmaier, 1968. 29.vi.2017, cord. Inorski (54044’15”N & 43008’53”E), Remark: The species is confirmed in MSNR. coll. G.B. Semishin. General distribution: Russia (the European part Ancistrocerus ichneumonideus (Ratzeburg, 1844) from the northwest till Crimea and northern Caucasus, Literature: Ruchin & Antropov 2016. Urals, Siberia, Amur region, Khabarovsk Krai and General distribution: Russia (the European part Primorsky Krai, and Sakhalin), Europe, Turkey, Caucasus, from the northwest to the northern Caucasus, Urals, Afghanistan, India, Kazakhstan, Kyrgyzstan, Mongolia, Altai, and eastern Siberia), Europe, Turkey, Caucasus, northeastern China, Japan (Hokkaido and Honshu), and Kazakhstan, and Mongolia (Fateryga 2017). North America (Fateryga 2017). Biology: Nests in existing cavities (in the stems Biology: Nests in existing cavities (in the stems of plants, on clay slopes, and in old nests of other

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Hymenoptera). Prey: caterpillars of various Lepidoptera, (Hokkaido) (Fateryga 2017). less often false larvae of Tenthredinidae and larvae of Biology: Nests in existing cavities (in the stems Curculionidae and Chrysomelidae. of plants, on clay slopes, and in old nests of other Hymenoptera). Prey: caterpillars of various Lepidoptera, Ancistrocerus nigricornis (Curtis, 1826) less often false larvae of sawflies (Tenthredinidae) and Literature: Redikortsev 1938 (as Odynerus callosus larvae of snout beetles (Curculionidae) and leaf beetles Thoms.); Plavilshchikov 1964 (as Odynerus callosus (Chrysomelidae). Predators: emerald wasps Chrysis Thms.); Mokrousov et al. 2009, 2013; Ruchin et al. 2009; ignita (Linnaeus, 1758), C. Impressa Schenck, 1856, and Ruchin & Antropov 2014. C. longula Abeille de Perrin, 1879 (Chrysididae). Material: RM: 1 female, 23.viii.2015, MSNR, quar. 387 (54045’43”N & 43015’46”E); 1 female, 10.v.2015, Ancistrocerus parietum (Linnaeus, 1758) quar. 447 (54043’14”N & 43013’39”E); 12.v.2016, Literature: Plavilshchikov 1964 (as Odynerus 1 female, quar. 408 (54044’57”N & 43011’47”E); 2 parietum L.); Ruchin et al. 2009; Mokrousov et al. 2009. female, 28.iv.2016, 6.viii.2017, quar. 420 (54045’29”N & General distribution: Russia (the European part from 43024’19”E); 1 female, 24.vii.2016, quar. 398 (54046’22”N the northwest till Crimea and northern Caucasus, Urals, & 43025’48”E); 2 female, 2 male, 23.viii.2016, quar. 85 Siberia, Amur region, Khabarovsk Krai and Primorsky (54053’29”N & 43035’00”E); 1 male, 10.viii.2016, quar. Krai, and Sakhalin), Europe, northern Africa, Turkey, 440 (54043’56”N & 43013’15”E); 3 female, 21.v.2016, Caucasus, Kazakhstan, ? central Asia, Iran, Mongolia, 21.v.2017, cord. Inorski (54044’15”N & 43008’53”E); and northern China. Acclimatized in North America as a 1 female, 9.v.2015, cord. Podrubnyi (54047’51”N & result of accidental importation (Fateryga 2017). 43008’48”E); 1 female, 18.v.2014, cord. Polyanski Biology: Nests in existing cavities (in the stems (54046’36”N & 43028’47”E); 1 female, 23.vii.2016, of plants, on clay slopes, and in old nests of other Elniki Distr., Novoyamskaya Sloboda (54044’24”N & Hymenoptera). Prey: caterpillars of various Lepidoptera, 43042’59”E). less often false larvae of sawflies (Tenthredinidae) General distribution: Russia (the European part and larvae of snout beetles (Curculionidae) and leaf from the northwest till Crimea and northern Caucasus, beetles (Chrysomelidae). Predators: emerald wasps Urals, Siberia, Khabarovsk Krai and Primorsky Krai, and Chrysis borealis Paukkunen, Ødegaard & Soon, 2015, Sakhalin), Europe, northern Africa, Kazakhstan, central C. fulgida Linnaeus, 1761, C. Ignita (Linnaeus, 1758), C. Asia, Iran, India, Mongolia, northeastern China, and ruddii Shuckard, 1837, C. sexdentata Christ, 1791, and Japan (Hokkaido and Honshu) (Fateryga 2017). Pseudochrysis neglecta (Shuckard, 1836) (Chrysididae). Biology: Nests in existing cavities (in the stems of plants, on clay slopes, and in old nests of other Ancistrocerus trifasciatus (Müller, 1776) Hymenoptera). Prey: caterpillars of various Lepidoptera, Literature: Redikortsev 1938 (as Odynerus less often false larvae of sawflies (Tenthredinidae) and trifasciatus F.); Plavilshchikov 1964 (as Odynerus larvae of snout beetles (Curculionidae) and leaf beetles trifasciatus F.); Ruchin et al. 2009; Mokrousov et al. (Chrysomelidae). Predators: emerald wasps Chrysis 2013; Ruchin & Antropov 2014. ignita (Linnaeus, 1758), C. Schencki Linsenmaier, 1968, Material: RM: 1 female, 15.viii.2014, MSNR, quar. and C. terminata Dahlbom, 1854 (Chrysididae). 420 (54045’29”N & 43024’19”E); 1 female, 12.vi.2016, quar. 413 (54045’12”N & 43016’54”E); 1 female, Ancistrocerus parietinus (Linnaeus, 1761) 1.vii.2016, quar. 329 (54046’36”N & 43010’39”E); 1 Literature: Mokrousov et al. 2013. female, 15.viii.2014, cord. Taratinski (54044’43”N & Material: RM: 11 female, 28.vii.2015, MSNR, quar. 79 43005’14”E); 1 female, 4.vii.2017, cord. Podrubnyi (54053’11”N & 43029’18”E); 1 female, 13.vii.2017, cord. (54047’51”N & 43008’48”E); 1 male, 18.vi.2017, cord. Inorski (54044’15”N & 43008’53”E), coll. G.B. Semishin; 4 Drozhdenovski (54044’31”N & 43017’24”E); NNR: 1 female, 12–16.v.2014, 27.v.2014, Pushta (54042’50”N & ex., 22.vii.2017, Voznesenskoe Distr., Alamasovo 43013’32”E). (54054’03”N & 43008’56”E), coll. P.V. Cherenkov (Image General distribution: Russia (the European part from 4). the northwest till Crimea and northern Caucasus, Urals, General distribution: Russia (the European part Siberia, Amur region, Khabarovsk Krai and Primorsky from the northwest till Crimea and northern Caucasus, Krai, and Sakhalin), Europe, northern Africa, Turkey, Urals, Siberia, Amur region, Kamchatka, Khabarovsk Krai Caucasus, Kazakhstan, Iran, Mongolia, Korea, and Japan and Primorsky Krai, Sakhalin, and Kuril Islands), Europe,

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Turkey, Caucasus, Kazakhstan, Kyrgyzstan, Mongolia, others), less often false larvae of sawflies (). northeastern China, Korea, and Japan (Hokkaido and Predators: emerald wasps Chrysis impressa Schenck, Honshu) (Fateryga 2017). 1856, C. inaequalis Dahlbom, 1845, and C. splendidula Biology: Nests in existing cavities (in the stems Rossi, 1790 (Chrysididae). of plants, on clay slopes, and in old nests of other Remark: The species is confirmed in MSNR. Hymenoptera). Prey: caterpillars of various Lepidoptera, less often false larvae of sawflies (Tenthredinidae) and Eumenes coronatus (Panzer, 1799) larvae of snout beetles (Curculionidae) and leaf beetles Literature: Mokrousov et al. 2013; Ruchin & (Chrysomelidae). Predators: emerald wasps Chrysis Antropov 2014. angustula Schenck, 1856, C. ignita (Linnaeus, 1758), C. Material: RM: 1 female, 22.viii.2015, MSNR, quar. impressa Schenck, 1856, C. mediata Linsenmaier, 1951, 342 (54046’21”N & 43023’23”E); 1 female, 4.viii.2017, C. schencki Linsenmaier, 1968, C. solida Haupt, 1957, quar. 413 (54045’12”N & 43016’54”E); 1 female, and C. subcoriacea Linsenmaier, 1959 (Chrysididae). 13.vii.2017, quar. 347 (54047’34”N & 43028’31”E); 1 female, 4.viii.2017, cord. Inorski (54044’15”N & Discoelius zonalis (Panzer, 1801) 43008’53”E), coll. G.B. Semishin; 1 male, 7.viii.2017, Literature: Redikortsev 1938 (as Discoelius naryshkini cord. Pavlovski (54045’15”N & 43024’03”E), coll. G.B. F. Моr.); Plavilshchikov 1964 (as Discoelius naryshkini F. Semishin; 1 female, 20.vii.2017, Pushta (54042’50”N & Моr.); Ruchin & Antropov 2016. 43013’32”E). General distribution: Russia (the European part General distribution: Russia (the European part from from the centre till Crimea and northern Caucasus, the northwest till Crimea and northern Caucasus, Urals, Urals, Siberia, Primorsky Krai, and Kuril Islands), Europe, Altai, eastern Siberia, Amur region, Khabarovsk Krai and Caucasus, Turkey, Iran, Mongolia, central and eastern Primorsky Krai, and Kuril Islands), Europe, Israel, Turkey, China, Korea, and Japan (Hokkaido, Honshu, Shikoku, Caucasus, Iran, Kazakhstan, central Asia, Mongolia, Kyushu, and Ryukyu) (Fateryga 2017). China, Korea, and Japan (Kyushu) (Fateryga 2017). Biology: Nests in the abandoned tunnels of the Biology: Nests of clay in the shape of a pot on branches beetle larvae in the wood. Prey: small caterpillars of bushes, stones, and walls of buildings, sometimes (Cochylidae, Pyralidae, Torticidae, Gelechiidae, in groups. Prey: small caterpillars (Geometridae, Eucosmidae, and Pyraustidae), less often false larvae Tortricidae, Pyralidae, Alucitidae, and Pyraustidae), less of sawflies (Pamphilidae). Predators: emerald wasps often false larvae of sawflies (Cimbicidae). Predators: Chrysis brevitarsis Thomson, 1870 and C. equestris emerald wasps Hedychrum rutilans Dahlbom, 1854 and Dahlbom, 1854 (Chrysididae). Chrysis ruddii Shuckard, 1837 (Chrysididae). Eumenes coarctatus (Linnaeus, 1758) Literature: Ruchin & Antropov 2016. Eumenes pedunculatus (Panzer, 1799) Material: RM: 1 female, 19.vii.2016, MSNR, quar. 330 Literature: Mokrousov et al. 2009, 2013; Feoktistov (54046’44”N & 43011’09”E); 1 male, 24.vii.2016, quar. 2011; Ruchin & Antropov 2014. 398 (54046’22”N & 43025’48”E); 1 male, 10.viii.2016, Material: RM: 1 female, 24.viii.2017, MSNR, quar. 80 0 0 quar. 427 (54044’30”N & 43012’49”E); 1 female, (54 53’11”N & 43 30’22”E); 1 male, 20.viii.2016, quar. 0 0 15.vi.2017, Temnikov Distr., Tyuveevo (54039’45”N & 417 (54 45’23”N & 43 20’56”E); 1 male, 13.viii.2016, 0 0 43014’43”E). quar. 446 (54 43’13”N & 43 13’23”E); 1 male, 0 0 General distribution: Russia (the European part from 6.viii.2017, quar. 419 (54 45’39”N & 43 22’50”E); 1 male, the northwest till Crimea and northern Caucasus, Urals, 2.vii.2017, cord. Inorski (54044’15”N & 43008’53”E); Siberia, Amur and Magadan regions, and Khabarovsk 1 male, 21.v.2017, Temnikov Distr., Russkoe Karaevo Krai and Primorsky Krai), Europe, northern Africa, Israel, (54040’55”N, 43013’52”E); 1 male, 15.vi.2017, Tyuveevo Jordan, Cyprus, Turkey, Syria, Iran, Kazakhstan, central (54039’45”N & 43014’43”E); 3 male, 23.vii.2016, Asia, Mongolia, China, and Japan (Honshu, Shikoku, and 22.vii.2017, Elniki Distr., Malye Mordovskie Poshaty Kyushu) (Fateryga 2017). (54041’00”N & 43043’50”E). Biology: Nests of clay in the shape of a pot on branches General distribution: Russia (the European part of bushes, stones, and walls of buildings, sometimes from the northwest to the centre and the east, Urals, in groups. Prey: small caterpillars (Geometridae, Siberia, Primorsky Krai, and Sakhalin), Europe, Turkey, Tortricidae, Pyralidae, Alucitidae, Pyraustidae, and Kazakhstan, Mongolia, and southwestern China

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(Fateryga 2017). Iran, Kazakhstan, and Mongolia (Fateryga 2017). Biology: Nests of clay in the shape of a pot on branches Biology: Nests in hollow stems of herbaceous plants of bushes, stones, and walls of buildings, sometimes or in branches of bushes with a soft core. Prey: larvae in groups. Prey: small caterpillars (Geometridae, of snout beetles Phytonomus (Curculionidae) and, Tortricidae, Pyralidae, Alucitidae, and Pyraustidae), less probably, catterpillars of Microlepidoptera. Predators: often false larvae of sawflies (Cimbicidae). emerald wasps Chrysis ignita (Linnaeus, 1758), C. indigotea Dufour & Perris, 1840, C. rutilans Olivier, 1791, ** Euodynerus notatus (Jurine, 1807) C. splendidula Rossi, 1790, and Pseudochrysis neglecta Material: RM: 1 female, 18.vi.2017, MSNR, cord. (Shuckard, 1836) (Chrysididae). Drozhdenovski (54044’31”N & 43017’24”E). General distribution: Russia (the European part Microdynerus parvulus (Herrich-Schäffer 1838) from the northwest till Crimea and northern Caucasus, Literature: Mokrousov et al. 2013. Urals, Altai, eastern Siberia, and also, perhaps, the Far General distribution: Russia (the European part from East (Amur and Magadan regions, Khabarovsk Krai and the northwest till Crimea and northern Caucasus, Urals, Primorsky Krai, and Sakhalin), Europe, northern Africa, and Altai), Europe, Caucasus, Turkey, Iran, Kazakhstan, Caucasus, Turkey, Iran, Kazakhstan, Mongolia, and, and Uzbekistan (Fateryga 2017). possibly, northern China (Fateryga 2017). Biology: Nests in hollow stems of herbaceous plants Biology: Nests in hollow stems of plants, tunnels of or in branches of bushes with a soft core. Prey: small bark beetles, and in old nests of Hymenoptera (including larvae of snout beetles (Curculionidae). Predators: in the ground). Prey: catterpillars of Tortricidae, Pyralidae, emerald wasps Chrysis gracillima Förster, 1853 Gelechiidae, Pyraustidae, and of other families, less (Chrysididae). often larvae of leaf beetles (Chrysomelidae). Predators: emerald wasps Chrysis graelsii Guérin-Méneville, 1842, Odynerus reniformis (Gmelin, 1790) C. pseudobrevitarsis Linsenmaier, 1951, C. sexdentata Literature: Mokrousov et al. 2009. Christ, 1791, C. solida Haupt, 1957, and C. zetterstedti Material: RM: 1 male, 18.vi.2017, MSNR, cord. Dahlbom, 1845 (Chrysididae). Drozhdenovski (54044’31”N & 43017’24”E); 1 female, 30.vi.2015, Temnikov Distr., Nizhnie Borki (54039’24”N Euodynerus quadrifasciatus (Fabricius, 1793) & 43028’31”E); 1 male, 13.viii.2015, Elniki Distr., Malye Literature: Ruchin & Antropov 2016. Mordovskie Poshaty (54041’00”N & 43043’50”E); NNR: 1 General distribution: Russia (the European part male, 4.vii.2017, Voznesenskoe Distr., Zarya (54050’08”N from the northwest till Crimea and northern Caucasus, & 43007’08”E). Urals, Siberia, Amur and Magadan regions, Khabarovsk General distribution: Russia (the European part Krai and Primorsky Krai, Sakhalin, and Kuril Islands), from the northwest to the centre and the east, Urals, Europe, northern Africa, Caucasus, Turkey, Kazakhstan, and Siberia), Europe, northern Africa, Israel, Turkey, and Mongolia, northeastern China, Korea, and Japan Kazakhstan (Fateryga 2017). (Hokkaido and Honshu) (Fateryga 2017). Biology: Nests in the soil, with a long curved Biology: Nests in hollow stems of plants, tunnels tube of clay lumps in front of the entrance, both on of bark beetles, and in old nests of Hymenoptera vertical and horizontal surfaces. Prey: larvae of snout (including in the ground). Prey: catterpillars of beetles Phytonomus (Curculionidae) and leaf beetles Tortricidae, Pyralidae, Gelechiidae, Pyraustidae, and (Chrysomelidae) and small caterpillars Microlepidoptera. of other families, less often larvae of leaf beetles Predators: emerald wasps Chrysis mediadentata (Chrysomelidae). Predators: emerald wasps Chrysis Linsenmaier, 1951, C. ruddii Shuckard, 1837, C. viridula graelsii Guérin-Méneville, 1842 and C. pseudobrevitarsis Linnaeus, 1761, and Pseudochrysis neglecta (Shuckard, Linsenmaier, 1951 (Chrysididae). 1836) (Chrysididae). Remark: The species is confirmed in MSNR. Gymnomerus laevipes (Shuckard, 1837) Literature: Ruchin & Antropov 2016. Odynerus simillimus F. Morawitz, 1867 General distribution: Russia (the European part Literature: Ruchin & Antropov 2014. from the northwest till Crimea and northern Caucasus, General distribution: Russia (the European part from Urals, Siberia, Amur region, Primorsky Krai, and the northwest till Crimea and northern Caucasus, Urals, Sakhalin), Europe, northern Africa, Caucasus, Turkey, and Siberia), west and east of Europe, Israel, Caucasus,

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Turkey, Kazakhstan, Turkmenistan, and Mongolia F. and O. bifasciatus L.); Plavilshchikov 1964 (as (Fateryga 2017). Discoelius sinuatus F., Odynerus bifasciatus L. and Biology: Nests in the soil, with a long curved tube of Odynerus diunatus F.); Ruchin et al. 2009; Mokrousov clay lumps in front of the entrance. Prey: larvae of snout et al. 2009, 2013; Ruchin et al. 2009; Feoktistov 2011 beetles (Curculionidae). (as Symmorphus mutinensis Bald.); Ruchin & Antropov 2014. * Stenodynerus bluethgeni van der Vecht, 1971 Material: RM: 1 female, 24.vii.2016, MSNR, quar. Material: RM: 1 male, 4.viii.2013, MSNR, quar. 330 398 (54046’22”N & 43025’48”E); 1 female, 6.viii.2017, (54046’44”N & 43011’09”E). quar. 397 (54046’17”N & 43025’44”E); 1 male, 15.vi.2014, General distribution: Russia (the European part from quar. 408 (54044’57”N & 43011’47”E); 1 male, 13.vi.2016, the northwest till Crimea and northern Caucasus, Urals, quar. 435 (54043’41”N & 43007’59”E); 1 female, 1 male, Novosibirsk region, Krasnoyarsk Krai, and Irkutsk region), 9.vi.2014, 16.vi.2014, Pushta (54042’50”N & 43013’32”E); south of Europe, Caucasus, Turkey, Iran, Kazakhstan, 1 male, 23.vii.2016, Elniki Distr., Cherlyaj (54045’53”N & Turkmenistan, and northern China (Fateryga 2017). 43040’58”E). General distribution: Russia (the European part * Stenodynerus picticrus (Thomson, 1874) from the northwest till Crimea and northern Caucasus, Material: RM: 1 female, 9.vi.2016, MSNR, quar. 331 Urals, Siberia, Amur region, Khabarovsk Krai and (54046’43”N & 43012’37”E). Primorsky Krai, and Sakhalin), Europe, Turkey, Caucasus, General distribution: Russia (the European part Kazakhstan, central Asia, Mongolia, China, Korea, and from the northwest to the centre, Urals, Siberia, Amur Japan (Hokkaido and Honshu) (Fateryga 2017). and Magadan regions, Kamchatka, Khabarovsk Krai Biology: Nests in a variety of existing cavities (galls, and Primorsky Krai, and Sakhalin), Europe, Turkey, hollow stems of plants, and in the old nests of other Kazakhstan, and Mongolia (Fateryga 2017). Hymenoptera in soil and wood). Prey: caterpillars Biology: Nests in a variety of existing cavities of Lithocolletidae, Tischeriidae, Stigmellidae, (galls, hollow stems of plants, and in the old tunnels of Cosmopterygidae, and Heliozelidae. Preys mostly beetles in the wood). Prey: caterpillars of Tortricidae, on larvae of Phratora species, sometimes catching Psychidae, Blastobasidae, Eucosmidae, Gelechiidae, Linaeidea and Plagiodera as well (Budrienė 2003). Lithocolletidae, and Pyralidae, less often larvae of leaf Predators: emerald wasps Schenck, beetles (Chrysomelidae). 1856, C. fulgida Linnaeus, 1761, and C. ignita (Linnaeus, 1758) (Chrysididae). Stenodynerus xanthomelas (Herrich-Schäffer, 1839) Literature: Ruchin & Antropov 2014, 2016. ** Symmorphus connexus (Curtis, 1826) Material: RM: 1 female, 17.vi.2016, MSNR, quar. 330 Material: RM: 1 male, 13.vi.2016, MSNR, quar. 421 (54046’44”N & 43011’09”E); 1 female, 12.vi.2016, quar. (54043’52’’N, 43007’06”E). 413 (54045’12”N & 43016’54”E); 1 male, 12.vi.2016, quar. General distribution: Russia (the north and the 430 (54044’34”N & 43016’00”E); 1 female, 15.vii.2017, centre of the European part and Altai), Europe, Caucasus, quar. 324 (54046’28”N & 43005’40”E). Turkey, and Kazakhstan (Fateryga 2017). General distribution: Russia (the European part Biology: Nests in a variety of existing cavities (galls, from the centre till Crimea and northern Caucasus and hollow stems of plants, and in the old nests of other Altai), south of Europe, Caucasus, Turkey, Iran, and Hymenoptera in soil and wood). Prey: caterpillars northeastern China (Fateryga 2017). of Lithocolletidae, Tischeriidae, Stigmellidae, Biology: Nests in a variety of existing cavities Cosmopterygidae, Heliozelidae, or larvae of snout (galls, hollow stems of plants, and in the old tunnels of beetles (Cucculionidae) and leaf beetles (Chrysomelidae). beetles in the wood). Prey: caterpillars of Tortricidae, Predators: emerald wasps Chrysis angustula Schenck, Psychidae, Blastobasidae, Eucosmidae, Gelechiidae, 1856 (Chrysididae). Lithocolletidae, and Pyralidae, less often larvae of leaf beetles (Chrysomelidae). Symmorphus crassicornis (Panzer, 1798) Remark: The species is confirmed in MSNR. Literature: Mokrousov et al. 2009, 2013; Feoktistov 2011; Ruchin & Antropov 2014. Symmorphus bifasciatus (Linnaeus, 1761) Material: RM: 1 male, 28.vii.2015, MSNR, quar. 82 Literature: Redikortsev 1938 (as Odynerus sinuatus (54053’12”N & 43032’46”E); 1 female, 27.v.2014, Pushta

13214 Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov

(54042’50”N & 43013’32”E). (Chrysomelidae). General distribution: Russia (the European part from the northwest till Crimea and northern Caucasus, Urals, Symmorphus murarius (Linnaeus, 1758) and Siberia), Europe, Turkey, Caucasus, Kazakhstan, Literature: Feoktistov 2011. central Asia, Iran, Mongolia, India, ? northeastern China Material: RM: 2 female, 25.v.2015, MSNR, quar. 0 0 (Fateryga 2017). 276 (54 47’45”N & 43 10’28”E); 1 female, 11.vii.2015, quar. 301 (54047’19”N & 43010’36”E); 1 female, 1 male, Biology: Nests in a variety of existing cavities (galls, 21.vi.2015, quar. 360 (54046’13”N & 43013’31”E); 4 hollow stems of plants, and in the old nests of other male, 21.vi.2015, quar. 381 (54045’17”N & 43009’52”E); 1 Hymenoptera in soil and wood). Prey: caterpillars male, 31.v.2015, quar. 398 (54046’22”N & 43025’48”E); 1 of Lithocolletidae, Tischeriidae, Stigmellidae, female, 9.vi.2016, quar. 331 (54046’43”N & 43012’37”E); Cosmopterygidae, Heliozelidae, or larvae of snout 1 female, 18.vii.2016, quar. 383 (54045’36”N & beetles (Curculionidae) and leaf beetles (Chrysomelidae) 43011’48”E). (Budrienė 2003). Predators: emerald wasps Chrysis General distribution: Russia (the European part from fulgida Linnaeus, 1761, C. iris Christ, 1791, and C. longula the northwest till Crimea and northern Caucasus, Urals, Abeille de Perrin, 1879 (Chrysididae). Tomsk region, eastern Siberia, and Primorsky Krai), Europe, northern Africa, Turkey, Caucasus, Kazakhstan, * Symmorphus fuscipes (Herrich-Schäffer, 1838) central Asia, Iran, and Korea (Fateryga 2017). Material: RM: 2 female, 14.vi.2016, MSNR, quar. 384 Biology: Nests in a variety of existing cavities (galls, (54045’33”N & 43012’44”E); 1 female, 11.vi.2016, quar. hollow stems of plants, and in the old nests of other 368 (54046’37”N & 43021’45”E); 1 female, 12.vi.2016, Hymenoptera in soil and wood). Prey: caterpillars quar. 430 (54044’34”N & 43016’00”E); 2 male, 9.vi.2016, of Lithocolletidae, Tischeriidae, Stigmellidae, quar. 358 (54046’05”N & 43011’44”E); 1 male, 18.vii.2016, Cosmopterygidae, and Heliozelidae, or larvae of snout beetles (Curculionidae) and leaf beetles (Chrysomelidae) quar. 383 (54045’36”N & 43011’48”E). (Budrienė 2003). Predators: Chrysis fulgida Linnaeus, General distribution: Russia (the European part 1761, C. iris Christ, 1791, and C. longula Abeille de from the northwest to the centre, Urals, Siberia, Amur Perrin, 1879 (Chrysididae). and Magadan regions, Primorsky Krai, and Sakhalin), Remark: The species is confirmed in MSNR. south of Europe, Azurbayjan, Kazakhstan, Mongolia, and northeastern China (Fateryga 2017). Subfamily Polistinae Biology: Nests in a variety of existing cavities (galls, * Polistes albellus Giordani Soika, 1976 hollow stems of plants, and in the old nests of other Literature: Ruchin et al. 2009 (as Polistes nimpha Hymenoptera in soil and wood). Prey: caterpillars (Christ, 1791), part.); Mokrousov et al. 2013 (as Polistes of Lithocolletidae, Tischeriidae, Stigmellidae, bischoffi Weyrauch, 1937); Ruchin & Antropov 2014 (as Cosmopterygidae, Heliozelidae, or (more often) larvae Polistes biglumis (Linnaeus, 1758)). of snout beetles (Curculionidae) and leaf beetles Material: RM: 1 female, 29.v.2016, MSNR, quar. 142 (Chrysomelidae). (54051’12”N & 43009’21”E); 1 female, 15.vii.2017, quar. 324 (54046’28”N & 43005’40”E); 1 female, 12.ix.2017, 0 0 Symmorphus gracilis (Brullé, 1832) quar. 115 (54 51’46”N & 43 09’26”E); 1 female, 0 Literature: Ruchin & Antropov 2016. 12.vii.2016, Temnikov Distr., Alkaevo (54 35’49”N & 43022’35”E); 1 female, 22.vii.2017, Elniki Distr., Novye General distribution: Russia (the European part Shaly (54042’13”N & 43038’25”E); 1 female, 22.vii.2017, from the centre till Crimea and northern Caucasus), Malye Mordovskie Poshaty (54041’00”N & 43043’50”E); Europe, Israel, Lebanon, Turkey, Caucasus, Kazakhstan, NNR: 1 female, 4.vii.2017, Voznesenskoe Distr., Zarya Turkmenistan, Tajikistan, and Iran (Fateryga 2017). (54050’08”N & 43007’08”E). Biology: Nests in a variety of existing cavities (galls, General distribution: Russia (the European part from hollow stems of plants, and in the old nests of other the centre to the south, Urals, Siberia, and ? Primorsky Hymenoptera in soil and wood). Prey: caterpillars Krai), south of Europe, Kazakhstan, and Mongolia of Lithocolletidae, Tischeriidae, Stigmellidae, (Antropov 2017a). Cosmopterygidae, Heliozelidae, or (more often) larvae Biology: Nests in the shape of a horizontal or inclined of snout beetles (Curculionidae) and leaf beetles open bottom comb structure on a single column on the

Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 13215 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov stems of plants or under stones. Prey: small caterpillars 1 female, 19.vi.2016, cord. Taratinski (54044’43”N & of Geometridae and Tortricidae. 43005’14”E); 3 female, 1.ix.2014, 19.v.2016, 21.v.2016, Pushta (54042’50”N & 43013’32”E); 1 female, 17.v.2014, Polistes dominula (Christ, 1791) Temnikov Distr., Tretyakovo (54031’44”N & 43013’18”E); Literature: Plavilshchikov 1964 (as Pollistes gallicus 3 male, 5.viii.2015, 15.viii.2015, Alkaevo (54035’49”N L.); Ruchin et al. 2009 (as Polistes dominulus (Christ, & 43022’35”E); 1 female, 1 male, 24.vii.2016, Polyanki 1791)); Ruchin & Antropov 2014. (54044’27”N & 43029’04”E); 5 female, 1 male, Material: NNR: 1 ex., 28.v.2017, Sarov (54055’08” N 19.vii.2015, 23.vii.2016, 22.vii.2017, Elniki Distr., Malye 43017’40”E), coll. P.V. Cherenkov. Mordovskie Poshaty (54041’00”N & 43043’50”E); 1 General distribution: Russia (the European part male, 23.vii.2016, Cherlyaj (54045’53”N & 43040’58”E); from the northwest till Crimea and northern Caucasus, NNR: 1 ex., 23.ix.2017, Voznesenskoe Distr., Alamasovo Urals, and Altai), Europe, northern Africa, Israel, (54054’03”N & 43008’56”E), coll. P.V. Cherenkov (Image Jordan, Turkey, Georgia, Armenia, Kazakhstan, central 4). Asia, Iran, Afghanistan, Pakistan, India, Mongolia, and General distribution: Russia (the European part northwestern China. Acclimatized in the Canary Islands, from the northwest till Crimea and the northern North and South America, and Australia (Antropov Caucasus, Urals, Siberia, Amur region, and Khabarovsk 2017a). Krai), Europe, northern Africa, Israel, Jordan, Turkey, Biology: Nests in the shape of a horizontal or inclined Georgia, Azerbaijan, Armenia, Kazakhstan, Kyrgyzstan, open bottom comb structure on a single column on the Iraq, Iran, Pakistan, India, Mongolia, and northeastern stems of plants or in shelters, on several columns. Prey: China (Antropov 2017a). caterpillars of Geometridae and Tortricidae (Hughes et Biology: Nests in the shape of a horizontal or inclined al. 2003). open bottom comb structure on a single column on the stems of plants or in shelters, on several columns (Kozyra Polistes nimpha (Christ, 1791) & Baraniak 2016). Prey: caterpillars of Geometridae and Literature: Redikortsev 1938 (Polistes opinabilis Tortricidae. Kohl); Plavilshchikov 1964 (Pollistes opinabilis Kohl); Ruchin et al. 2009; Mokrousov et al. 2013 (Pollistes Subfamily Vespinae nimphus (Christ, 1791)); Ruchin & Antropov 2014. Vespa crabro Linnaeus, 1758 Material: RM: 1 female, 28.vii.2015, MSNR, quar. 79 Literature: Redikortsev 1938; Plavilshchikov 1964; (54053’11”N & 43029’18”E); 2 male, 16.viii.2015, quar. Ruchin et al. 2009; Mokrousov et al. 2013. 324 (54046’28”N & 43005’40”E); 1 female, 5.vii.2015, Material: RM: 1 ex., 24.v.2015, quar. 448 quar. 403(54044’40”N & 43006’57”E); 1 female, 7.ix.2015, (54042’37”N & 43012’29”E); 1 ex., 10.viii.2016, quar. quar. 430 (54044’34”N & 43016’00”E); 1 female, 427 (54044’30”N & 43012’49”E); 1 ex., 28.vii.2015, quar. 20.iv.2014, quar. 434 (54045’04”N & 43019’02”E); 1 79 (54053’11”N & 43029’18”E); 1 ex., 16.viii.2015, quar. female, 24.v.2015, quar. 448 (54042’37”N & 43012’29”E); 324 (54046’28”N & 43005’40”E); 1 ex., 19.vii.2016, cord. 1 female, 29.v.2016, quar. 35 (54053’19”N & Podrubnyi (54047’51”N & 43008’48”E); 2 ex., 28.vi.2015, 43011’11”E); 1 male, 23.viii.2016, quar. 59 (54054’14”N & 11.v.2015, cord. Inorski (54044’15”N & 43008’53”E); 43034’57”E); 1 male, 23.viii.2016, quar. 85 (54053’29”N & 1 ex., 19.vi.2016, cord. Taratinski (54044’43”N & 43035’00”E); 1 female, 18.vi.2016, quar. 197 (54050’05”N 43005’14”E); 1 ex., 18.vi.2017, cord. Drozhdenovski & 43009’09”E); 1 female, 18.vi.2016, quar. 275 (54044’31”N & 43017’24”E); 3 ex., 27.v.2014, 1.ix.2014, (54047’46”N & 43008’34”E); 1 female, 5.vii.2016, quar. Pushta (54042’50”N & 43013’32”E); 1 ex., 17.v.2014, 324 (54046’28”N & 43005’40”E); 1 female, 11.vi.2016, Temnikov Distr., Tretyakovo (54031’44”N & 43013’18”E); quar. 368 (54046’37”N & 43021’45”E); 1 female, 1 ex., 15.viii.2015, Alkaevo (54035’49”N & 43022’35”E); 1 19.vi.2016, quar. 401 (54044’30”N & 43005’15”E); 2 ex., 22.vii.2017, Elniki Distr., Malye Mordovskie Poshaty male, 20.viii.2016, quar. 417 (54045’23”N & 43020’56”E); (54041’00”N & 43043’50”E); 1 ex., 23.vii.2016, Cherlyaj 1 male, 3.ix.2016, quar. 421 (54043’52’’N, 43007’06”E); 1 (54045’53”N & 43040’58”E); NNR: 1 ex., 4.vii.2017, male, 10.viii.2016, quar. 427 (54044’30”N & 43012’49”E); Voznesenskoe Distr., Zarya (54050’08”N & 43007’08”E); 3 female, 26.v.2016, 13.viii.2016, quar. 446 (54043’13”N 1 ex., 23.viii.2017, Diveevo Distr., Satis (54055’25”N & 43013’23”E); 4 female, 6.ix.2014, 28.vi.2015, 11.v.2015, & 43013’24”E); 1 ex., 6.viii.2014, Sarov (54055’08” N cord. Inorski (54044’15”N & 43008’53”E); 2 female, 43017’40”E), coll. P.V. Cherenkov (Image 3). 19.vii.2016, cord. Podrubnyi (54047’51”N & 43008’48”E); General distribution: Russia (the European part from

13216 Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov the northwest till Crimea and northern Caucasus, Urals, quar. 86 (54053’38”N & 43035’59”E); 1 female, Siberia, Amur region, Khabarovsk Krai and Primorsky 4.v.2016, quar. 380 (54045’06”N & 43008’34”E); 1 Krai, and Sakhalin), Europe, Turkey, Azerbaijan, Georgia, female, 2 male, 15.v.2016, 18.vii.2016, quar. 383 Kazakhstan, Turkmenistan, Iran, Mongolia, China, Korea, (54045’36”N & 43011’48”E); 1 female, 13.vi.2016, and Japan (Hokkaido and Honshu). Acclimatized in quar. 435 (54043’41”N & 43007’59”E); 1 female, 2 North America and Guatemala as a result of accidental male, 10.vi.2016, 24.vii.2016, quar. 440 (54043’56”N & importation (Antropov 2017a). 43013’15”E); 3 female, 1.viii.2017, quar. 111; 5.vii.2015, Biology: As a rule, nests are in shelters, in hollows of 23.viii.2017, 4 female, cord. Inorski (54044’15”N & trees and in attics. The shell of the nest is relatively fragile 43008’53”E), coll. G. B. Semishin; 2 female, 11.vii.2015, since the material is wood fibres of decayed stumps and 20.v.2017, cord. Podrubnyi (54047’51”N & 43008’48”E); trunks. In closed hollows, the outer shell can be absent. 1 female, 9.vi.2016, cord. Novenkovski (54055’53”N & Prey: various insects from large Diptera to honey bees 43025’18”E); 3 female, 22.vi.2014, 8.v.2015, 5.vii.2016, and male . The family development cycle is Pushta (54042’50”N & 43013’32”E); NNR: 2 female, long, from May to October. 1.viii.2017, Pervomaisk Distr., 6 km southwest of Lesozavod (54054’32”N & 43038’11”E). Dolichovespula media (Retzius, 1783) General distribution: Russia (the European part Literature: Redikortsev 1938 (as Vespa media Deg.); from the northwest till Crimea and northern Caucasus, Plavilshchikov 1964 (as Vespula media Deg.); Ruchin et Urals, Siberia, Amur and Magadan regions, Chukotka, al. 2009; Mokrousov et al. 2013. Kamchatka, Khabarovsk Krai and Primorsky Krai, Material: RM: 1 female, 31.vi.2015, MSNR, quar. 345 Sakhalin, and Kuril Islands), Europe, Turkey, Kazakhstan, (54047’13”N & 43026’30”E); 1 male, 18.vii.2016, quar. Iran, Mongolia, southwestern and northern China, 383 (54045’36”N & 43011’48”E); 1 male, 6.viii.2017, quar. Korea, and Japan (Hokkaido and Honshu) (Antropov 397 (54046’17”N & 43025’44”E); 1 female, 22.vii.2017, 2017a). Elniki Distr., Malye Mordovskie Poshaty (54041’00”N & Biology: Quite common, sometimes abundant 43043’50”E). species. Nests have gray smooth outer shell, openly General distribution: Russia (the European part from hanging on branches of trees or bushes or in shelters the northwest till Crimea and northern Caucasus, Urals, (in household outbuildings and attics). An immature Siberia, Amur region, ? Kamchatka, Khabarovsk Krai nest does not have an entry tube. Prey: mostly various and Primorsky Krai, Sakhalin, and Kuril Islands), Europe, dipterans. The family development cycle is short, from Morocco, Turkey, Georgia, Syria, Kazakhstan, Mongolia, May to August. northeastern China, Korea, and Japan (Hokkaido and Honshu). Accidentally imported introducted to New Dolichovespula sylvestris (Scopoli, 1763) Zealand (Antropov 2017a). Literature: Plavilshchikov 1964 (as Vespula silvestris Biology: Nests have thick gray outer shell, openly Scop.); Mokrousov et al. 2009, 2013. hanging in the middle or upper tiers of branches of Material: RM: 2 female, 18.vi.2016, MSNR, quar. trees and shrubs or under the eaves of buildings. An 197 (54050’05”N & 43009’09”E); 1 male, 24.vii.2016, immature nest has a more or less long entry tube. Prey: quar. 398 (54046’22”N & 43025’48”E); 1 female, 1 male, mostly various Diptera. The family development cycle is 19.vii.2015, 23.vii.2016, Elniki Distr., Malye Mordovskie short, from the end of May to August. Poshaty (54041’00”N & 43043’50”E); 1 male, 23.vii.2016, Novoyamskaya Sloboda (54044’24”N & 43042’59”E); 1 Dolichovespula saxonica (Fabricius, 1793) male, 23.vii.2016, Cherlyaj (54045’53”N & 43040’58”E). Literature: Mokrousov et al. 2009, 2013; Feoktistov General distribution: Russia (the European part 2011; Ruchin & Antropov 2014. from the northwest till Crimea and northern Caucasus, Material: RM: 1 female, 1 male, 28.vii.2015, Urals, Siberia, Amur region, and Khabarovsk Krai and MSNR, quar. 79 (54053’11”N & 43029’18”E); 1 female, Primorsky Krai), Europe, Morocco, Turkey, Armenia, 28.vii.2015, quar. 82 (54053’12”N & 43032’46”E); 1 Georgia, Kazakhstan, Kyrgyzstan, Uzbekistan, Syria, female, 4.vii.2015, quar. 288 (54048’27”N & 43022’03”E); Iran, Afghanistan, Pakistan, India, Mongolia, and China 2 female, 10.v.2015, quar. 366 (54046’37”N & (Antropov 2017a). 43019’49”E); 1 female, 21.vi.2015, quar. 381 (54045’17”N Biology: Nests have gray loose outer shell, openly & 43009’52”E); 2 female, 9.v.2015, 5.vii.2015, quar. hanging in the lower tier of branches of bushes or stems 403 (54044’40”N & 43006’57”E); 1 female, 12.vii.2014, of grass. An immature nest does not have an entry tube.

Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 13217 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov

Prey: mostly various dipterans. The family development & 43013’32”E); 1 female, 21.v.2017, Temnikov Distr., cycle is short, from May to early August (Edwards & Russkoe Karaevo (54040’55”N, 43013’52”E); 1 female, Telfer 2002). The species prefers coniferous forests 23.vii.2016, Elniki Distr., Cherlyaj (54045’53”N & (Dvořák 2007). 43040’58”E). General distribution: Russia (the European part Vespula germanica (Fabricius, 1793) from the northwest till Crimea and northern Caucasus, Literature: Plavilshchikov 1964 (as Pseudovespa Urals, Siberia, Amur and Magadan regions, Chukotka, germanica F.); Mokrousov et al. 2009; Ruchin & Kamchatka, Khabarovsk Krai and Primorsky Krai, Antropov 2014. Sakhalin, and Kuril Islands), Europe, Turkey, Georgia, Material: RM: 1 female, 12.ix.2017, MSNR, quar. Kazakhstan, Uzbekistan, Kyrgyzstan, Turkmenistan, Iran, 115 (54051’46”N & 43009’26”E); 2 female, 9.ix.2016, Mongolia, , China, Korea, and Japan (Hokkaido, 12.ix.2017, cord. Plotomoika (54053’36”N & 43009’41”E); Honshu, and Shikoku) (Antropov 2017a). 3 female, 17.vii.2017, 5.ix.2017, 29.ix.2017, Pushta Biology: Round nests have a gray loose outer shell, (54042’50”N & 43013’32”E); 1 female, 23.vii.2016, Elniki located mainly in cavities under the roots of trees or in Distr., Cherlyaj (54045’53”N & 43040’58”E); NNR: 1 ex., abandoned mammal burrows. An immature nest does 31.viii.2014, 4.x.2014, Sarov (54055’08” N 43017’40”E), not have an entrance tube. Prey: various Diptera. The coll. P.V. Cherenkov (Image 3). family development cycle is short, from May till late General distribution: Russia (the European part August. from the northwest till Crimea and northern Caucasus, Urals, Siberia, Amur region, Khabarovsk Krai and Vespula vulgaris (Linnaeus, 1758) Primorsky Krai, and Sakhalin), Europe, northern Africa, Literature: Redikortsev 1938; Plavilshchikov 1964 (as Israel, Jordan, Lebanon, Turkey, Armenia, Georgia, Pseudovespa vulgaris L.); Ruchin et al. 2009; Mokrousov Kazakhstan, Uzbekistan, Turkmenistan, Tajikistan, Syria, et al. 2009. Iraq, Iran, Afghanistan, Pakistan, India, Mongolia, China, Material: RM: 1 female, 28.vii.2015, MSNR, quar. and Korea. The species was accidentally introduced 82 (54053’12”N & 43032’46”E); 1 female, 20.vii.2015, with commercial cargo to Iceland, Madeira, Azores and quar. 301 (54047’19”N & 43010’36”E); 1 female, Canary Islands, South Africa, Australia, Tasmania, New 22.viii.2015, quar. 342 (54046’21”N & 43023’23”E); 3 Zealand, Fr. Ascension, Fr. Norfolk, Hawaii, and northern female, 24.viii.2014, 10.v.2015, quar. 368 (54046’37”N & (USA and Canada) and southern (Argentina and Chile) 43021’45”E); 1 female, 22.viii.2015, quar. 396; 1 female, America. 26.v.2015, quar. 447 (54043’14”N & 43013’39”E); 1 Biology: Nests have a gray undulating outer shell, female, 12.vii.2014, quar. 86 (54053’38”N & 43035’59”E); with short posts on the outside, located mainly in 1 female, 15.v.2015, quar. 420 (54045’29”N & underground cavities (abandoned mammalian burrows) 43024’19”E); 1 female, 24.vii.2016, quar. 398 (54046’22”N or under buildings. The species is quite common in the & 43025’48”E); 1 female, 13.viii.2016, quar. 446 reserve. As in other parts of the range (Dvořák 2007), (54043’13”N & 43013’23”E); 1 female, 21.v.2017, quar. it prefers glades, clearings, meadows, and forest edges. 421 (54043’52’’N, 43007’06”E); 1 female, 13.vii.2014, An immature nest does not have an entrance tube. cord. Novenkovski (54055’53”N & 43025’18”E); 1 female, Prey: various Diptera. The family development cycle is 16.v.2014, cord. Srednayay Melnitsa (54054’09”N & long, from May to October (Edwards & Telfer 2002). 43013’53”E), coll. L.V. Egorov; 2 female, 11.v.2015, 28.vi.2015, cord. Inorski (54044’15”N & 43008’53”E); 2 Vespula rufa (Linnaeus, 1758) female, 16.vii.2016, 31.v.2016, Pushta (54042’50”N & Literature: Ruchin et al. 2009; Feoktistov 2011 (as 43013’32”E); 1 female, 23.vii.2016, Elniki Distr., Cherlyaj Paravespula rufa L.); Ruchin & Antropov 2014. (54045’53”N & 43040’58”E); NNR: 1 ex., 6.viii.2014, Sarov Material: RM: 1 female, 20.vi.2015, MSNR, quar. 345 (54055’08” N 43017’40”E), coll. P.V. Cherenkov; 1 ex., (54047’13”N & 43026’30”E); 1 female, 23.viii.2015, quar. 7.ix.2014, Voznesenskoe Distr., Alamasovo (54054’03”N 360 (54046’13”N & 43013’31”E); 1 female, 10.v.2014, & 43008’56”E), coll. P.V. Cherenkov. quar. 408 (54044’57”N & 43011’47”E); 1 female, General distribution: Russia (the European 23.viii.2016, quar. 85 (54053’29”N & 43035’00”E); 1 part from the northwest till Crimea and northern female, 3.vii.2016, quar. 283 (54048’08”N & 43017’28”E); Caucasus, Urals, Siberia, Amur and Magadan regions, 1 female, 10.viii.2016, quar. 427 (54044’30”N & Chukotka, Kamchatka, Khabarovsk Krai and Primorsky 43012’49”E); 1 female, 31.vii.2015, Pushta (54042’50”N Krai, Sakhalin, and Kuril Islands), Europe, Turkey,

13218 Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov

Georgia, Kazakhstan, Kyrgyzstan, Iran, India, Mongolia, Primorsky Krai), Europe, Turkey, Kazakhstan, Iran, southwestern and northeastern China, Korea, and Mongolia, northern and southwestern China, and North Japan (Hokkaido and Honshu). As a result of accidental America (Danilov 2017). importation, it acclimatized in Iceland, Australia, New Biology: Nests in dense sandy soil, often in groups. Zealand, Mauri, and Hawaii (Antropov 2017a). Prey: caterpillars of Geometridae and , and Biology: Nests have an ochreish undulating outer extremely rarely false larvae of Tentredinidae (Kazenas shell, without any posts on the outside, located mainly 2001). Predators: fliesMetopia argyrocephala (Meigen, in underground cavities (abandoned mammalian 1824) (Sarcophagidae) and twisted-wing parasites burrows) or in wall cavities and under buildings. An Paraxenos sphecidarum (Dufour, 1837) (Stylopidae). immature nest does not have an entrance tube. Prey: various Diptera. The family development cycle is long, sabulosa (Linnaeus, 1758) from May to October (Edwards & Telfer 2002). It is the Literature: Plavilshchikov 1964; Ruchin et al. 2009; most widespread and numerous species of social wasps Mokrousov et al. 2009, 2013; Ruchin & Antropov 2014. in the reserve. Material: RM: 2 female, 23.viii.2015, MSNR, quar. 360 (54046’13”N & 43013’31”E); 1 female, 17.viii.2014, Superfamily Apoidea quar. 397 (54046’17”N & 43025’44”E); 1 male, 20.vi.2015, Family Sphecidae quar. 345 (54047’13”N & 43026’30”E); 1 male, 21.vi.2015, Subfamily Ammophilinae quar. 381 (54045’17”N & 43009’52”E); 1 male, 5.vii.2015, Ammophila campestris (Latreille, 1809) quar. 403 (54044’40”N & 43006’57”E); 1 male, 5.vii.2015, Literature: Mokrousov et al. 2013. quar. 405; 1 male, 12.vii.2014, quar. 86 (54053’38”N & General distribution: Russia (the European part 43035’59”E); 1 female, 23.viii.2016, quar. 85 (54053’29”N from the northwest till Crimea and northern Caucasus, & 43035’00”E); 1 female, 18.vii.2016, quar. 331 Urals, Siberia, Amur region, and Primorsky Krai), Europe, (54046’43”N & 43012’37”E); 1 male, 1.vii.2016, quar. 379 northern Africa, Turkey, Kazakhstan, central Asia, (54045’17”N & 43007’51”E); 1 male, 7.vii.2016, quar. 386 Mongolia, northern China, and Korea (Danilov 2017). (54045’40”N & 43014’48”E); 1 female, 6.viii.2017, quar. Biology: Nests in clay or dense sandy soil. Prey: 420 (54045’29”N & 43024’19”E); 2 female, 12.ix.2017, caterpillars of Geometridae, Pieridae, and Noctuidae quar. 60 (54053’08”N & 43009’57”E); 1 male, 13.vii.2017, and false larvae of sawflies (Tentredinidae). Predators: quar. 347 (54047’34”N & 43028’31”E); 1 male, flies Metopia (Sarcophagidae). 13.vii.2017, quar. 357 (54046’07”N & 43010’56”E); 2 male, 13.vii.2017, quar. 319 (54048’09”N & 43028’25”E); Ammophila pubescens Curtis, 1836 1 female, 24.vii.2016, Temnikov Distr., Polyanki Literature: Mokrousov et al. 2013; Ruchin & (54044’27”N & 43029’04”E); 1 female, 23.vii.2016, Elniki Antropov 2014. Distr., Cherlyaj (54045’53”N & 43040’58”E); 1 male, Material: RM: 1 female, 28.vii.2015, MSNR, quar. 79 22.vii.2017, Novoyamskaya Sloboda (54044’24”N & (54053’11”N & 43029’18”E); 1 female, 12.vii.2014, quar. 43042’59”E). 86 (54053’38”N & 43035’59”E); 1 female, 11.vii.2015, General distribution: Russia (the European part from quar. 142 (54051’12”N & 43009’21”E); 1 female, the northwest till Crimea and northern Caucasus, Urals, 18.vi.2016, quar. 115 (54051’46”N & 43009’26”E); 1 Siberia, Magadan region, Kamchatka, and Khabarovsk male, 13.vii.2017, quar. 319 (54048’09”N & 43028’25”E); Krai and Primorsky Krai), Europe, northern Africa, 4.vi.2017, 1 female, cord. Inorski, coll. G.B. Semishin, Turkey, Syria, Iran, Kazakhstan, central Asia, Mongolia, 2.vii.2017, 1 male, cord. Inorski (54044’15”N & and northern China (Danilov 2017). 43008’53”E); 1 female, 22.vii.2017, Elniki Distr., Biology: Nests in dense sandy soil, sometimes on Novoyamskaya Sloboda (54044’24”N & 43042’59”E); steep slopes with coarse-grained soil. Prey: large naked 2 female, 1 male, 31.v.2014, 22.vii.2017, Malye caterpillars of Geometridae, Limantriidae, Pieridae, Mordovskie Poshaty (54041’00”N & 43043’50”E); NNR: Notodontidae, and Noctuidae (Casiraghi et al. 2001b; 3 female, 4.vii.2017, 16.vii.2017, Voznesenskoe Distr., Kazenas 2001). Predators: some Ichneumonidae Zarya (54050’08”N & 43007’08”E); 1 female, 16.vii.2017, (Casiraghi et al. 2001a). Svobodnyi (54050’47”N & 43007’48”E). General distribution: Russia (the European part ? Ammophila terminata F. Smith, 1856 from the northwest till Crimea and northern Caucasus, Literature: Feoktistov 2011. Urals, Siberia, Amur region, and Khabarovsk Krai and General distribution: Russia (the European part

Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 13219 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov from the centre till Crimea, Urals, and Siberia), Europe, stictica (Meigen, 1830) (Sarcophagidae), and twisted- northern Africa, Turkey, Kazakhstan, central Asia, wing parasites Paraxenos sphecidarum (Dufour, 1837) Mongolia, and northern China (Yildirim et al. 2016; (Stylopidae). There are cases of stealing prey from other Danilov 2017). nests. Biology: Nests in dense sandy soil. Prey: caterpillars of geometer moths (Geometridae). ? tydei (Le Guillou, 1841) Remark: Confirmation of the species in Mordovia Literature: Plavilshchikov 1964 (as Psammophila requires more material. tydei Guill.). General distribution: Russia (the European part from Podalonia affinis (W. Kirby, 1798) the centre till Crimea and northern Caucasus), south Literature: Redikortsev 1938; Plavilshchikov 1964 (as of Europe, Africa, Israel, Jordan, Turkey, Kazakhstan, Psammophila affinis Kby.); Mokrousov et al. 2013. central Asia, Syria, Iraq, Iran, Afghanistan, India, Pakistan, General distribution: Russia (the European part from Mongolia, and northwest of China (Murai & Amr 2011). the northwest till Crimea and northern Caucasus, Urals, As a result of accidental importation, acclimatized in the Siberia, Amur region, and Khabarovsk Krai and Primorsky Canary Islands and Australia (Danilov 2017). Krai), north and east of Europe, northern Africa, Israel, Remark: Confirmation of the species in Mordovia Jordan, Turkey, Caucasus, Kazakhstan, central Asia, Iran, requires more material. Mongolia, southwestern and northern China, and Korea (Danilov 2017). Subfamily Sceliphrinae Biology: Nests in coarse-grained or dense sandy soil. Sceliphron deforme (F. Smith, 1856) Prey: caterpillar of owlet moths Agrotis (Noctuidae) Literature: Ćetković et al. 2011; Mokrousov et al. living in the ground. Predators: some Ichneumonidae 2013; Ruchin & Antropov 2014. (Casiraghi et al. 2001a). Material: RM: 1 male, 9 female, 7.vii.2014, 8.vii.2014, 26.vii.2017, 6.viii.2017, MSNR, cord. Inorski (54044’15”N Podalonia hirsuta (Scopoli, 1763) & 43008’53”E), coll. G.B. Semishin; 1 female, 1 male, Literature: Redikortsev 1938 (as Ammophila hirsuta 16.vii.2017, cord. Podrubnyi (54047’51”N & 43008’48”E); Scop.); Plavilshchikov 1964 (as Psammophila hirsuta 2 female, 1 male, 4.viii.2017, 7.viii.2017, cord. Pavlovski Scop.); Ruchin et al. 2009; Mokrousov et al. 2009. (54045’15”N & 43024’03”E), coll. G.B. Semishin; 2 female, Material: RM: 1 female, 11.vii.2015, MSNR, quar. 301 1 male, 20.v.2014, 13.vii.2014, 31.v.2014, Pushta (54047’19”N & 43010’36”E); 1 female, 14.v.2016, quar. (54042’50”N & 43013’32”E). 278 (54047’51”N & 43012’27”E); 1 female, 15.v.2016, General distribution: Russia (the European quar. 276 (54047’45”N & 43010’28”E); 2 female, part from the centre to northern Caucasus, Urals, 23.viii.2016, quar. 85 (54053’29”N & 43035’00”E); 1 Novosibirsk region, Krasnoyarsk Krai, Amur region, male, 18.vii.2016, quar. 278 (54047’51”N & 43012’27”E); and Khabarovsk Krai and Primorsky Krai), southeastern 1 male, 18.vii.2016, quar. 303 (54047’17”N & Europe, Kazakhstan, Uzbekistan, Kyrgyzstan, Tajikistan, 43012’32”E); 1 male, 18.vii.2016, quar. 331 (54046’43”N Mongolia, China, Korea, Japan (Hokkaido, Honshu, & 43012’37”E); 1 female, 14.v.2017, quar. 327; 1 female, Shikoku, Kyushu, and Zyukyu), India, and southeastern 15.v.2017, quar. 302 (54047’15”N & 43011’26”E); NNR: Asia (Danilov 2017). 1 ex., 16.vii.2017, Voznesenskoe Distr., Alamasovo Biology: Nests are placed in various shelters, built (54054’03”N & 43008’56”E), coll. P.V. Cherenkov. of clay cells molded into a single layer without covering General distribution: Russia (the European part from by a common shell, but surrounded by a roll of clay. In the northwest till Crimea and northern Caucasus, Urals, the reserve it is found in residential and commercial and Siberia), Europe, northern Africa, Israel, Lebanon, premises. This species often makes nests over windows, Turkey, Caucasus, Kazakhstan, central Asia, Iran, in door jambs, under eaves, and under roofs. Prey: Afghanistan, Pakistan, India, Mongolia, and western and spiders of medium size, usually the most common and northern China (Danilov 2017). available in the habitat (Gepp 2003). Biology: Hibernate in the imago stage. Nests in coarse-grained or dense sandy soil. Prey: caterpillars, Sceliphron destillatorium (Illiger, 1807) mainly Cucullia and Agrotis (Noctuidae), sometimes Literature: Mokrousov et al. 2013; Ruchin & Epinephele (Satyridae). Predators: parasitic flies Antropov 2014. Miltogramma punctatum Meigen, 1824, Taxigramma Material: RM: 1 female, 3.vii.2014, MSNR, cord.

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Inorski (54044’15”N & 43008’53”E), coll. G.B. Semishin; Elniki Distr., Malye Mordovskie Poshaty (54041’00”N & 2 female, 19.vii.2015, 22.vii.2017, Elniki Distr., Novye 43043’50”E). Shaly (54042’13”N & 43038’25”E). General distribution: Russia (the European part General distribution: Russia (the European part from from the northwest till Crimea and northern Caucasus, the centre till Crimea and northern Caucasus, Urals, Urals, Siberia, Amur region, and Khabarovsk Krai and Altai, and Transbaikalia), south of Europe, northern Primorsky Krai), Europe, northern Africa, Israel, Yemen, Africa, Israel, Turkey, Kazakhstan, central Africa, Syria, Oman, Turkey, Abkhazia, Kazakhstan, Uzbekistan, Iran, Afghanistan, Mongolia, and China (Danilov 2017). Turkmenistan, Tajikistan, Iraq, Iran, Afghanistan, India, Biology: Nests of clay cells molded into a few layers, Mongolia, northern China, and Korea (Murai & Amr covered by a common shell, in open spaces or in shelters. 2011; Nemkov 2017a). Prey: spiders of medium size, usually the most common Biology: Nests in dense soil. Prey: larvae of the late and available in the habitat (Gepp 2003). stages of heteropterans Cydnidae and Pentatomidae. Predators: emerald wasps Hedychridium roseum (Rossi, Subfamily Sphecinae 1790), H. rossicum Gussakovskij, 1948, and Holopyga Tribe Pryonichini generosa (Förster, 1853) (Chrysididae). Prionyx nudatus (Kohl, 1885). Literature: Ruchin & Antropov 2014. Dryudella lineata Mocsáry, 1879 General distribution: Russia (the European part from Literature: Mokrousov et al. 2013. the centre till Crimea and northern Caucasus, Urals, General distribution: Russia (from the centre to Novosibirsk region, Altai, and Tuva), south and east of the south of the European part), east of Europe, and Europe, northern Africa, Turkey, Kazakhstan, central Kazakhstan (Nemkov 2017a). Asia, Iran, Afghanistan, and northern China (Danilov 2017). ** Dryudella stigma (Panzer, 1809) Biology: Nests in dense sandy soil. Prey: imago or Literature: Mokrousov et al. 2013. older larva of locusts (Acrididae). Material: RM: 1 female, 18.vii.2016, MSNR, quar. 303 (54047’17”N & 43012’32”E). Tribe Sphecini General distribution: Russia (the European part Sphex funerarius Gussakovskij, 1934 from the northwest to the centre and east, Altai, eastern Material: NNR: 2 ex., 5.viii.2014, 11.viii.2014, Sarov Siberia, and Primorsky Krai), Europe, and Kazakhstan (54055’08” N 43017’40”E), coll. P.V. Cherenkov (Image (Nemkov 2017a). 4). Biology: Nests in dense sandy soil. Prey: nymphs General distribution: Russia (the European part of Heteroptera, mostly Lygaeidae. Predators: emerald from the centre till Crimea and northern Caucasus, wasps Hedychridium cupreum (Dahlbom, 1845), H. Urals, Siberia, and Khabarovsk Krai), Europe, northern femoratum (Dahlbom, 1854), and Holopyga metallica Africa, Israel, Turkey, Georgia, Kazakhstan, central Asia, (Dahlbom, 1854) (Chrysididae). Syria, Iran, Afghanistan, Mongolia, and northern China Remark: The species is confirmed in MSNR. (Danilov 2017). Biology: Nests in dense sandy or clay soil. Prey: Subfamily Bembicinae grasshoppers Decticus verrucivorus (Linnaeus, 1758), Tribe Alysontini Phaneroptera phalcata (Poda, 1761), Platycleis grisea Didineis lunicornis (Fabricius, 1798) (= Didineis (Fabricius, 1781), P. intermedia (Audinet-Serville, 1838), kozhevnikovi Kokujev, 1906) Tettigonia albifrons (Fabricius, 1775), and T. viridissima Literature: Mokrousov et al. 2013. (Linnaeus, 1758) (Tettigonidae). General distribution: Russia (centre, south, and east of the European part and Urals), Europe, northern Family Crabronidae Africa, and Abkhazia (Nemkov 2017b). Subfamily Astatinae Biology: Nests in dense soil. Prey: Astata boops (Schrank, 1781) and, probably, Heteroptera. Literature: Mokrousov et al. 2013. Material: RM: 1 female, 13.vii.2015, MSNR, quar. 434 Tribe (54045’04”N & 43019’02”E); 1 male, 28.vii.2015, quar. Bembix rostrata (Linnaeus, 1758) 79 (54053’11”N & 43029’18”E); 1 female, 19.vii.2015, Literature: Redikortsev 1938 (as Bembex rostrata L.);

Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 13221 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov

Plavilshchikov 1964 (as Bembex rostrata L.); Mokrousov 1 female, 13.vii.2017, quar. 347 (54047’34”N & et al. 2013; Ruchin & Antropov 2014. 43028’31”E); 1 female, 11.vii.2015, cord. Podrubnyi Material: RM: 1 male, 12.vii.2014, MSNR, quar. (54047’51”N & 43008’48”E). 86 (54053’38”N & 43035’59”E); 1 female, 24.viii.2017, General distribution: Russia (the European part quar. 347 (54047’34”N & 43028’31”E); 2 male, 8.vi.2014, from the north to the centre and the east, Urals, Altai, 28.vi.2015, cord. Inorski; 1 male, 27.vii.2017, cord. Krasnoyarsk Krai, and Irkutsk region), Europe, Turkey, Inorski (54044’15”N & 43008’53”E), coll. G.B. Semishin; and Kazakhstan (Nemkov 2017b). 1 male, 13.vii.2014, cord. Novenkovski (54055’53”N & Biology: Nests in dense (clay or gravel) soil. Prey: 43025’18”E); 1 male, 20.vi.2015, Temnikov Distr., Russkoe nymphs of leafhoppers, mainly Karaevo (54040’55”N, 43013’52”E); 1 female, 2.vii.2016, (). Kushki (54029’49” N 43016’11”E); 2 female, 23.vii.2016, 22.vii.2017, Elniki Distr., Malye Mordovskie Poshaty laticinctus (Lepeletier de Saint Fargeau, 1832) (54041’00”N & 43043’50”E); 1 female, 23.vii.2016, Literature: Redikortsev 1938 (as Gorytes laticinctus Novoyamskaya Sloboda (54044’24”N & 43042’59”E); Schr.); Plavilshchikov 1964 (as Gorytes laticinctus 1 male, 22.vii.2017, Novye Shaly (54042’13”N & Shuck.); Mokrousov et al. 2013; Ruchin & Antropov 43038’25”E); NNR: 1 male, 16.vii.2017, Voznesenskoe 2014. Distr., Svobodnyi (54050’47”N & 43007’48”E); 1 ex., Material: RM: 1 ex., 2.vi.2017, MSNR, cord. Inorski 8.viii.2014, Sarov (54055’08” N 43017’40”E), coll. P.V. (54044’15”N & 43008’53”E), coll. G.B. Semishin (Image Cherenkov (Images 3 & 4). 4). General distribution: Russia (the European part from General distribution: Russia (the European part from the northwest till Crimea and northern Caucasus, Urals, the northwest till Crimea and Altai), Europe, northern and Siberia), Europe, Turkey, Azerbaijan, Kazakhstan, Africa, Turkey, Kazakhstan, and Uzbekistan (Nemkov Uzbekistan, Turkmenistan, Tajikistan, Iran, Afghanistan, 2017b). Mongolia, and northern China (Nemkov 2017b). Biology: Nests in sandy soil. Prey: larvae and Biology: Nests in loose sandy soil. Prey: dipterans imagos of leafhoppers. Predators: digger wasps Nysson Tabanidae, Syrphidae, Muscidae, Calliphoridae, trimaculatus (Rossi, 1790) (Crabronidae). Sarcophagidae, and Antomyiidae. Predators: emerald wasps Parnopes grandior (Pallas, 1771), dipterans * Gorytes neglectus Handlirsch, 1895 Petrorossia hesperus Rossi, 1790 (Bombylidae), Metopia Material: RM: 1 female, 2.vi.2017, MSNR, cord. leucocephala (Rossi, 1790), Senotainia conica (Fallen, Inorski (54044’15”N & 43008’53”E), coll. G.B. Semishin. 1810), and Physocephala chrysorrhoea (Meigen, 1824) General distribution: Russi (the European part from (Sarcophagidae). the northwest to the centre, Siberia, Khabarovsk Krai and Primorsky Krai, and Sakhalin), northeast of Europe, * fargeii (Shuckard, 1837) Turkey, Kazakhstan, and Korea (Nemkov 2017b). Material: RM: 1 male, 31.v.2014, Elniki Distr., Malye Mordovskie Poshaty (54041’00”N & 43043’50”E). * Gorytes pleuripunctatus (A. Costa, 1859) General distribution: Russia (the European part Material: RM: 1 male, 22.vii.2017, Elniki Distr., from the north till Crimea, Urals, and Siberia), Europe, Novoyamskaya Sloboda (54044’24”N & 43042’59”E). Israel, Turkey, Armenia, Kazakhstan, Mongolia, and General distribution: Russia (the European part northeastern China (Nemkov 2017b). from the centre till Crimea and northern Caucasus and Biology: Nests in dense (clay, gravel, or coarse Urals), south of Europe, northern Africa, Israel, Turkey, skeletal) soil. Prey: nymphs of leafhoppers Jassidae. Georgia, Azerbaijan, Armenia, Kazakhstan, and Syria (Nemkov 2017b). Argogorytes mystaceus (Linnaeus, 1761) Literature: Mokrousov et al. 2009, 2013; Feoktistov Gorytes quadrifasciatus (Fabricius, 1804) 2011; Ruchin & Antropov 2014. Literature: Redikortsev 1938; Plavilshchikov 1964; Material: RM: 1 female, 12.vi.2016, MSNR, quar. 413 Mokrousov et al. 2013; Ruchin & Antropov 2014. (54045’12”N & 43016’54”E); 1 female, 12.vi.2016, quar. Material: RM: 3 male, 28.vii.2015, MSNR, quar. 82 430 (54044’34”N & 43016’00”E); 1 female, 13.vi.2016, (54053’12”N & 43032’46”E); 1 male, 11.vii.2015, quar. quar. 421 (54043’52’’N, 43007’06”E); 1 female, 115 (54051’46”N & 43009’26”E); 2 male, 11.vii.2015, 13.vi.2016, quar. 422 (54043’55”N & 43007’39”E); quar. 301 (54047’19”N & 43010’36”E); 1 male, 5.vii.2015,

13222 Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov quar. 405; 6 male, 18.vii.2016, quar. 331 (54046’43”N & imagos of leafhoppers. 43012’37”E); 2 male, 18.vii.2016, quar. 358 (54046’05”N & 43011’44”E); 1 male, 24.viii.2017, quar. 86 (54053’38”N Harpactus lunatus (Dahlbom, 1832) & 43035’59”E); 1 female, 11.viii.2014, Temnikov Distr., Literature: Mokrousov et al. 2013. Tatarskoe Karaevo (54042’17”N & 43013’49”E); 1 female, General distribution: Russia (the European part 24.vii.2016, Polyanki (54044’27”N & 43029’04”E); 1 male, from the north to the centre and the east, Altai, 23.vii.2016, Elniki Distr., Malye Mordovskie Poshaty Krasnoyarsk Krai, Irkutsk region, and Yakutia), Europe, (54041’00”N & 43043’50”E). and Kazakhstan (Nemkov 2017b). General distribution: Russia (the European part Biology: Nests in sandy soil. Prey: nymphs and imagos from the north till Crimea and northern Caucasus, Urals, of leafhoppers (Jassidae). Predators: emerald wasps Hedychridium cupreum (Dahlbom, 1845) (Chrysidae). Siberia, Amur and Magadan regions, Khabarovsk Krai and Primorsky Krai, and Sakhalin), Europe, northern Harpactus morawitzi Radoszkowski, 1884 Africa, Turkey, Kazakhstan, Kyrgyzstan, Mongolia, and Literature: Mokrousov et al. 2013. Korea (Nemkov 2017b). General distribution: Russia (the European part Biology: Nests in sandy or clay soil. Prey: nymphs and from the centre to northern Caucasus, Irkutsk region, Philaenus imagos of leafhoppers, mainly (Cercopidae). Buryatia, and Transbaikalia), south of Europe, Turkey, Predators: digger wasps Nysson maculosus (Gmelin, Kazakhstan, and Turkmenistan (Nemkov 2017b). 1790) and N. trimaculatus (Rossi, 1790) (Crabronidae). * Bembecinus hungaricus (Frivaldszky, 1876) Gorytes quinquecinctus (Fabricius, 1793) Material: RM: 1 female, 2.vii.2016, Temnikov Literature: Redikortsev 1938 (as Gorytes Distr., Kushki (54029’49” N 43016’11”E); NNR: 1 female, quadricinctus F.); Plavilshchikov 1964 (as Gorytes 16.vii.2017, Voznesenskoe Distr., Svobodnyi (54050’47”N quadricinctus F.); Mokrousov et al. 2013. & 43007’48”E). Material: RM: 1 male, 28.vii.2015, MSNR, quar. 82 General distribution: Russia (the European part from (54053’12”N & 43032’46”E); 1 male, 12.vii.2014, quar. the centre till Crimea and northern Caucasus), south of 86 (54053’38”N & 43035’59”E); 1 male, 11.vii.2015, Europe, Turkey, Abkhazia, and Kazakhstan (Nemkov quar. 115 (54051’46”N & 43009’26”E); 1 female, 3 male, 2012, 2017b). 20.vii.2014, quar. 276 (54047’45”N & 43010’28”E); 6 Biology: Nests in sandy soil, more often along the male, 20.vii.2014, 11.vii.2015, quar. 301 (54047’19”N banks of rivers and streams. Prey: larvae and imagos & 43010’36”E); 1 female, 27.vii.2014, quar. 384 of leafhoppers and jumping plant lice (Cicadellidae, (54045’33”N & 43012’44”E); 2 male, 13.vii.2015, quar. Psyllidae, Jassidae, and Fulgoridae). Predators: emerald 434 (54045’04”N & 43019’02”E); 1 male, 19.vii.2016, quar. wasps Spinolia unicolor (Dahlbom, 1831) (Chrysididae). 330 (54046’44”N & 43011’09”E); 2 female, 18.vii.2016, quar. 331 (54046’43”N & 43012’37”E); 1 male, 4.vii.2017, Bembecinus tridens (Fabricius, 1781) Literature: Mokrousov et al. 2013; Ruchin & quar. 357 (54046’07”N & 43010’56”E); 1 male, 8.vi.2014, Antropov 2014. cord. Inorski (54044’15”N & 43008’53”E); 1 female, Material: RM: 1 male, 2 female, 11.vii.2015, 15.viii.2014, cord. Taratinski (54044’43”N & 43005’14”E); 3.viii.2015, 18.vi.2016, MSNR, quar. 115 (54051’46”N 1 female, 23.vii.2016, Elniki Distr., Novoyamskaya & 43009’26”E); 1 female, 11.vii.2015, quar. 142 Sloboda (54044’24”N & 43042’59”E); 4 male, 23.vii.2016, (54051’12”N & 43009’21”E); 1 male, 28.vi.2015, cord. 0 22.vii.2017, Malye Mordovskie Poshaty (54 41’00”N & Inorski (54044’15”N & 43008’53”E); 1 female, 24.vi.2014, 0 0 43 43’50”E); 2 male, 23.vii.2016, Cherlyaj (54 45’53”N Pushta (54042’50”N & 43013’32”E); 1 female, 1 male, 0 & 43 40’58”E); NNR: 2 male, 4.viii.2017, Voznesenskoe 7.vii.2013, Temnikov Distr., Tarkhany; 2 male, 6.vii.2013, 0 0 Distr., Zarya (54 50’08”N & 43 07’08”E). Lavrentevo (54029’44”N & 43002’41”E); 1 female, General distribution: Russia (the European part 24.vii.2016, Polyanki (54044’27”N & 43029’04”E); 1 male, from the northwest till Crimea and northern Caucasus, 2.vii.2016, Kushki (54029’49” N 43016’11”E); 3 female, Urals, and Siberia), Europe, northern Africa, Turkey, 19.vii.2015, 22.vii.2017, Elniki Distr., Malye Mordovskie Azerbaijan, Kazakhstan, Uzbekistan, Kirghizia, Syria, and Poshaty (54041’00”N & 43043’50”E); 1 male, 22.vii.2017, Iran (Nemkov 2017b). Novye Shaly (54042’13”N & 43038’25”E). Biology: Nests in sandy soil. Prey: nymphs and General distribution: Russia (the European part from

Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 13223 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov the centre till Crimea and northern Caucasus, Urals, and northern Africa, Israel, Turkey, Abkhazia, and Kazakh- Altai), south of Europe, northern Africa, Israel, Jordan, stan (Nemkov 2017b). Lebanon, Oman, Turkey, Abkhazia, Georgia, Azerbaijan, Biology: Brood parasite of Gorytes, Harpactus, and Armenia, and Syria (Nemkov 2012, 2017b). Argogorytes (Crabronidae) [Blösch 2000]. Biology: Nests in loose sandy soil. Prey: larvae and imagos of leafhoppers (Fulgoridae, Cercopidae, Nysson maculosus (Gmelin, 1790) and Jassidae) and jumping plant lice (Psyllidae). Literature: Redikortsev 1938 (as Nysson maculatus Predators: velvet ants Nemka viduata (Pallas, 1773) and E.); Plavilshchikov 1964 (as Nysson maculatus F.); Mok- Smicromyrme nigriceps Nonveiller, 1959 (Mutillidae). rousov et al. 2013; Ruchin & Antropov 2016. General distribution: Russia (the European part ** Stizus perrisii Dufour, 1838 from the northwest till Crimea, Urals, Siberia, Amur re- Material: RM: 1 male, 4.vii.2017, MSNR, quar. 357 gion, Khabarovsk Krai and Primorsky Krai, and Sakhalin), (54046’07”N & 43010’56”E). Europe, northern Africa, Turkey, Georgia, Armenia, Ka- General distribution: Russia (the European part from zakhstan, Uzbekistan, Tajikistan, Mongolia, northern the northwest till Crimea and northern Caucasus, Urals, China, and Korea (Nemkov 2017b). Altai, Transbaikalia, and Primorsky Krai), Europe, Turkey, Biology: Brood parasite of digger wasps Gorytes Armenia, Kazakhstan, Uzbekistan, Kyrgyzstan, northern quadrifasciatus (Fabricius, 1804), Harpactus luna- China, Korea, and Japan (Honshu) (Nemkov 2017b). tus (Dahlbom, 1832), H. tumidus (Panzer, 1801), and Biology: Nests in dense clay or sandy soil. Prey: Lestiphorus bicinctus (Rossi, 1794). nymphs and imagos of grasshoppers () and locusts (Acrididae). (J. Forster, 1771) Remark: Confirmation of the species in Mordovia Literature: Ruchin et al. 2009; Mokrousov et al. requires more material. 2009; Feoktistov 2011; Mokrousov et al. 2013. Material: RM: 1 male, 21.vi.2015, MSNR, quar. 360 Brachystegus scalaris (Illiger, 1807) (54046’13”N & 43013’31”E); 1 female, 2 male, 29.v.2015, Literature: Mokrousov et al. 2013. 13.vi.2016, quar. 421 (54043’52’’N, 43007’06”E); 1 male, Material: RM: 2 female, 12.vii.2014, MSNR, quar. 86 26.v.2015, quar. 447 (54043’14”N & 43013’39”E); 2 male, (54053’38”N & 43035’59”E); 1 male, 13.vii.2014, cord. 5.vi.2015, quar. 448 (54042’37”N & 43012’29”E); 1 fe- Novenkovski (54055’53”N & 43025’18”E). male, 13.vi.2016, quar. 422 (54043’55”N & 43007’39”E); General distribution: Russia (the European part 2 male, 1 female, 13.vi.2016, quar. 435 (54043’41”N & from the centre till Crimea and northern Caucasus, and 43007’59”E); 1 male, 10.vi.2016, quar. 440 (54043’56”N Altai), south of Europe, northern Africa, Yemen, Turkey, & 43013’15”E); 1 male, 12.vi.2016, quar. 413 (54045’12”N Georgia, Kazakhstan, Uzbekistan, Kyrgyzstan, Tajikistan, & 43016’54”E); 1 male, 14.vi.2016, quar. 384 (54045’33”N and Iran (Nemkov 2017b). & 43012’44”E); 1 female, 15.vii.2017, quar. 324 Biology: Brood parasite of digger waspsTachytes (54046’28”N & 43005’40”E); 1 female, 8.vi.2014, cord. panzeri (Dufour, 1841) (Crabronidae) Inorski (54044’15”N & 43008’53”E). General distribution: Russia (the European part from Nysson dimidiatus Jurine, 1807 the northwest to northern Caucasus, Siberia, Amur re- Literature: Mokrousov et al. 2013. gion, Khabarovsk Krai and Primorsky Krai, Sakhalin, and General distribution: Russia (from the northwest to Kuril Islands), Europe, Turkey, Azerbaijan, Kazakhstan, northern Caucasus, Altai, Irkutsk region, Transbaikalia, Iran, and Japan (Honshu) (Nemkov 2017b). Amur and Magadan regions, and Sakhalin), Europe, Biology: Brood parasite of digger wasps Argogorytes Armenia, Kazakhstan, central Asia, and Mongolia fargeii (Shuckard, 1837), A. mystaceus (Linnaeus, 1761), (Nemkov 2017b). Gorytes laticinctus (Lepeletier de Saint Fargeau, 1832), Biology: Brood parasite of Harpactus and Bembecinus and G. quadrifasciatus (Fabricius, 1804). (Crabronidae) (Olberg 1959). Subfamily Nysson fulvipes A. Costa, 1859 Tribe Literature: Mokrousov et al. 2013. Entomognathus brevis (Vander Linden, 1829) General distribution: Russia (the European part Literature: Mokrousov et al. 2013. from the centre to northern Caucasus), south of Europe, General distribution: Russia (the European part

13224 Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov from the northwest till Crimea and northern Caucasus, Mongolia, and Korea (Antropov 2017b). Urals, Altai, Irkutsk region, Yakutia, Amur region, and Biology: Nests in the soil. Prey: various flies. Khabarovsk Krai and Primorsky Krai), Europe, northern Predators: emerald wasps Pseudomalus auratus Africa, Israel, Turkey, Georgia, Kazakhstan, Uzbekistan, (Linnaeus, 1758) and Trichrysis cyanea (Linnaeus, 1758) Turkmenistan, Tajikistan, Syria, Iran, Mongolia, northern (Chrysididae) and velvet ants Myrmosa atra Panzer, China, and Japan (Hokkaido and Honshu) (Antropov 1801 and Paramyrmosa brunnipes (Lepeletier, 1845) 2017b). (Mutillidae). Biology: Nests in the soil. Prey: leaf beetles of the Chaetocnema, less often Crepidodera, Crabro scutellatus (von Scheven, 1781) Cryptocephalus, Longitarsus, and Aphthoma Literature: Plavilshchikov 1964; Mokrousov et al. (Chrysomelidae). 2013. Material: RM: 1 female, 21.vi.2015, MSNR, quar. 408 Crabro cribrarius (Linnaeus, 1758) (54044’57”N & 43011’47”E); 1 female, 19.vi.2016, quar. Literature: Redikortsev 1938; Plavilshchikov 1964; 401 (54044’30”N & 43005’15”E); 1 male, 28.vi.2015, Mokrousov et al. 2013. cord. Inorski (54044’15”N & 43008’53”E); NNR: 2 female, Material: RM: 1 male, 11.vii.2015, MSNR, quar. 301 16.vii.2016, Voznesenskoe Distr., Svobodnyi (54050’47”N 0 0 (54 47’19”N & 43 10’36”E); 1 male, 21.vi.2015, quar. 384 & 43007’48”E). 0 0 (54 45’33”N & 43 12’44”E); 1 female, 18.vii.2016, quar. General distribution: Russia (the European part from 0 0 331 (54 46’43”N & 43 12’37”E); 2 male, 19.vii.2016, the northwest till Crimea and northern Caucasus, Urals, quar. 330 (54046’44”N & 43011’09”E); 1 female, Siberia, Amur region, Khabarovsk Krai and Primorsky 24.vii.2016, quar. 398 (54046’22”N & 43025’48”E); 2 Krai, and Sakhalin), Europe, Kazakhstan, Mongolia, and male, 4.vii.2017, quar. 357 (54046’07”N & 43010’56”E); eastern China (Antropov 2017b). 1 female, 11.vii.2015, cord. Podrubnyi (54047’51”N Biology: Nests in the soil, sometimes in burrows & 43008’48”E); 1 female, 23.vii.2016, Elniki Distr., of rodents. Prey: various flies (mainly Dolichopodidae Novoyamskaya Sloboda (54044’24”N & 43042’59”E); 1 (Diaphorus), sometimes Muscidae and Syrphidae). male, 23.vii.2016, Cherlyaj (54045’53”N & 43040’58”E). Predators: emerald wasps Chrysis fulgida Linnaeus, General distribution: Russia (the European part from 1761 (Chrysididae). the northwest till Crimea and northern Caucasus, Urals, Siberia, Amur region, Khabarovsk Krai and Primorsky Crossocerus annulipes (Lepeletier et Brullé, 1835) Krai, and Sakhalin), Europe, northern Africa, Turkey, Literature: Mokrousov et al. 2013. Kazakhstan, Iran, Mongolia, and Korea (Antropov 2017b). General distribution: Russia (the European part Biology: Nests are usually in the soil, rarely in rotten from the northwest till Crimea and northern Caucasus, stump wood. Prey: various flies. Predators: emerald Urals, Altai, Irkutsk region, Amur region, and Primorsky wasps Hedychridium coriaceum (Dahlbom, 1854) Krai), Europe, northern Africa, Israel, Turkey, Abkhazia, (Chrysididae). Kazakhstan, Uzbekistan, Kyrgyzstan, Mongolia, and North America (Antropov 2017b). Crabro peltarius (Schreber, 1784) Biology: Nests in abandoned tunnels of xylophagous Literature: Plavilshchikov 1964; Mokrousov et al. insects in wood or in hollow stems. Prey: dipterans. 2013. Material: RM: 1 female, 19.vi.2016, MSNR, quar. 401 Crossocerus assimilis (F. Smith, 1856) (54044’30”N & 43005’15”E); 1 female, 13.vii.2016, quar. Literature: Mokrousov et al. 2013. 319 (54048’09”N & 43028’25”E); 1 female, 3.vii.2016, General distribution: Russia (the European part quar. 338 (54047’06”N & 43019’40”E); NNR: 1 female, from the centre to the northern Caucasus, Urals, Altai, 1.viii.2017, Pervomaisk Distr., 6 km southwest of Irkutsk region, Transbaikalia, Amur region, Khabarovsk Lesozavod (54054’32”N & 43038’11”E). Krai and Primorsky Krai, Sakhalin, and Kuril Islands), General distribution: Russia (the European part from Europe, Turkey, Kazakhstan, Mongolia, Korea, and Japan the northwest till Crimea and northern Caucasus, Urals, (Honshu) (Antropov 2017b). Siberia, Amur region, Khabarovsk Krai and Primorsky Biology: Forest species. Nests in the wood of Krai, and Sakhalin), Europe, Turkey, Kazakhstan, deciduous tree species. Prey: small flies of the genus

Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 13225 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov

Phora (Phoridae). (Aphidae). Remark: Confirmation of the species in Mordovia Crossocerus barbipes (Dahlbom, 1845) requires more material. Literature: Mokrousov et al. 2013. General distribution: Russia (the European part Crossocerus leucostoma (Linnaeus, 1758) from the northwest to the centre and the east, Urals, Literature: Mokrousov et al. 2013; Ruchin & Siberia, Khabarovsk Krai and Primorsky Krai, Sakhalin, Antropov 2016. and Kuril Islands), Japan, Turkey, Kazakhstan, Mongolia, Material: RM: 1 female, 11.vi.2016, MSNR, quar. 368 southwestern China, Korea, Japan (Hokkaido, Honshu, (54046’37”N & 43021’45”E); 1 female, 11.vii.2017, quar. Yakushima, and Ryukyu), and North America (Antropov 408 (54044’57”N & 43011’47”E); 2 female, 31.v.2014, 2017b). Elniki Distr., Malye Mordovskie Poshaty (54041’00”N & Biology: Nests in wood. Prey: leafhoppersEmpoasca 43043’50”E). and Typhlocyba (Cicadellidae). General distribution: Russia (the European part from the northwest to the centre and east, Urals, Altai, Crossocerus cetratus (Shuckard, 1837) Irkutsk region, Magadan region, Kamchatka, Khabarovsk Literature: Mokrousov et al. 2013. Krai and Primorsky Krai, and Sakhalin), Europe, Turkey, Material: RM: 1 female, 24.vii.2016, MSNR, quar. Kazakhstan, Mongolia, Korea, Japan (Hokkaido and 0 0 399 (54 46’40”N & 43 27’27”E). Honshu), and North America (Antropov 2017b). General distribution: Russia (the European part Biology: Nests in abandoned tunnels of xylophagous from the northwest till Crimea and northern Caucasus, insects in dry wood. Prey: small dipterans. Predators: Urals, Altai, Irkutsk region, Primorsky Krai, Sakhalin, and ichneumons Tryphon signator Gravenhorst 1829 Kuril Islands), Europe, Turkey, Kazakhstan, Mongolia, (Ichneumonidae) and parasitic flies (Tachinidae). southwestern China, Korea, and Japan (Hokkaido and Honshu) (Antropov 2017b). Crossocerus megacephalus (Rossi, 1790) Biology: Forest species. Nests in abandoned tunnels Literature: Mokrousov et al. 2009. of xylophagous insects in wood and in branches of plants General distribution: Russia (the European with a soft core. Prey: small dipterans (Bibionidae, part from the northwest till Crimea and northern Ceratopogonidae, Chironomidae, Empididae, Miceto- Caucasus, Urals, Tomsk region, Altai, Irkutsk and Amur philidae, and Stratiomyiidae). regions, and Primorsky Krai), Europe, northern Africa, Turkey, Kazakhstan, Tajikistan, Iran, India, Mongolia, Crossocerus elongatulus (Vander Linden, 1829) northwestern China, and Japan (Hokkaido) (Antropov Literature: Mokrousov et al. 2013. General distribution: Russia (the European part from 2017b). the northwest till Crimea and northern Caucasus, Urals, Biology: Nests in abandoned tunnels of xylophagous Altai, Krasnoyarsk Krai, Irkutsk region, Primorsky Krai, insects in rotten wood and in branches with a soft core. Sakhalin, and Kuril Islands), Europe, northern Africa, Prey: small dipterans (Anthomyiidae, Chloropidae, Turkey, Azerbaijan, Kazakhstan, Kyrgyzstan, Tajikistan, Coelopidae, Dolichopodidae, Empididae, Lauxaniidae, Turkmenistan, Iran, Mongolia, and Japan (Hokkaido). Lonchaeidae, Muscidae, Rhagionidae, Sepsidae, Acclimatized in Canada, USA, and Argentina (Antropov Simuliidae, Stratiomyidae, Syrphidae, and Trypetidae). 2017b). Predators: chalcid wasps Habritys brevicornis (Ratzeburg, 1844) (), ichneumons Endasys Crossocerus exiguus (Vander Linden, 1829) analis (Thomson, 1883) (Ichneumonidae), and flies Literature: Mokrousov et al. 2013. Eustalomyia festiva (Zetterstedt, 1845) (Anthomyidae). General distribution: Russia (the European part Remark: Confirmation of the species in Mordovia from the northwest to the centre and the east, Urals, requires more material. Novosibirsk region, Altai, Irkutsk region, Yakutia, Transbaikalia, Amur region, and Khabarovsk Krai and ? Crossocerus ovalis Lepeletier de Saint Fargeau et Primorsky Krai), Europe, northern Africa, Kazakhstan, Brullé, 1835 Turkmenistan, Mongolia, northeastern China, and Korea Literature: Feoktistov 2011. (Antropov 2017b). General distribution: Russia (the European part from Biology: Nests in loose soil. Prey: various aphids the northwest to northern Caucasus, Urals, and Altai),

13226 Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov

Europe, Kazakhstan, and Mongolia (Antropov 2017b). (Sylvicola fenestralis (Scopoli, 1763)), Anthomyiidae, Biology: Nests in sandy or clay soil. Prey: mostly Chironomidae, Culicidae, Dolichopodidae, Ephydridae, small dipterans Tachydromia arrogans (Linnaeus, Helomyzidae, Lauxaniidae, Muscidae, Rhagionidae, and 1761) (Empididae), Platypalpus verralli (Collin, 1926) Syrphidae (Sphaerophoria scripta (Linnaeus, 1758)), (Hybotidae), Chironomidae or Chloropidae, sometimes sometimes small lepidopterans (Lepidoptera) and Heteroptera Plagiognathus chrysanthemi (Wolff, 1804) caddisflies (Trichoptera). (Miridae), and rarely small lepidopterans. Remark: Confirmation of the species in Mordovia * Crossocerus vagabundus (Panzer, 1798) requires more material. Material: RM: 1 female, 15.vi.2016, MSNR, quar. 429 (54044’31”N & 43014’59”E). Crossocerus palmipes (Linnaeus, 1767) General distribution: Russia (the European part from Literature: Mokrousov et al. 2013. the northwest till Crimea and northern Caucasus, Urals, General distribution: Russia (the European part Tomsk region, Altai, Krasnoyarsk Krai, Irkutsk region, from the northwest to the centre and the east, Urals, Transbaikalia, Amur region, Kamchatka, Khabarovsk Novosibirsk region, Altai, Krasnoyarsk Krai, Irkutsk Krai and Primorsky Krai, and Sakhalin), Europe, northern region, and Buryatia), Europe, northern Africa, Turkey, Africa, Turkey, Kazakhstan, Uzbekistan, India, Mongolia, Kazakhstan, and Mongolia (Antropov 2017b). China, Korea, and Japan (Hokkaido, Honshu, Shikoku, Biology: Nests in sandy soil. Prey: small dipterans and Kyushu) (Antropov 2017b). (Chloropidae, Dolichopodidae, Drosophilidae, Biology: Nests are linear or include several branches Lauxaniidae, Muscidae, and, probably, Empididae). in abandoned tunnels of xylophagous insects in wood Predators: velvet ants Myrmosa atra Panzer, 1801 or in hollow stems. Prey: dipterans (Chironomidae, (Mutillidae). Muscidae, Rhagionidae, Tabanidae, and Tipulidae), as an exeption leafroller moths (Tortricidae) and even worker Crossocerus podagricus (Vander Linden, 1829) ants Myrmica rubra (Linnaeus 1758) (Formicidae). Literature: Plavilshchikov 1964 (as Crabro podagricus Lind.). * Crossocerus wesmaeli (Vander Linden, 1829) General distribution: Russia (the European part from Material: RM: MSNR, quar. 401 (54044’30”N & the north till Crimea and northern Caucasus, Urals, Altai, 43005’15”E), 19.vi.2016, 1 female. Primorsky Krai, Sakhalin, and Kuril Islands), Europe, General distribution: Russia (the European part northern Africa, Turkey, Kazakhstan, Iran, Mongolia, from the northwest to northern Caucasus, Urals, Tomsk Korea, and Japan (Hokkaido and Honshu) (Antropov region, Altai, Irkutsk region, Amur region, Kamchatka, 2017b). Primorie Territory, Sakhalin, and Kuril Islands), Europe, Biology: Forest species. Nests in tunnels of northern Africa, Turkey, Kazakhstan, Uzbekistan, and xylophagous insects in wood. Prey: various small Tajikistan(Antropov 2017b). dipterans (Ceratopogonidae, Chironomidae, Empididae, Biology: Nests in dry soil, sometimes in abandoned Lycoriidae, Mycetophilidae, Phoridae, Scatopsidae, and tunnels of other insects. Prey: various small dipterans Sciaridae). (Agromyzidae, Ceratopogonidae, Chironomidae, Chloropidae, Dolichopodidae, Empididae, Ephydridae, Crossocerus quadrimaculatus (Fabricius, 1793) Lauxaniidae, Lonchaeidae, Muscidae, Simuliidae Literature: Ruchin & Antropov 2014. (Simulium equinum (Linnaeus, 1758), Tephritidae, and Material: RM: 1 female, 5.vii.2015, MSNR, quar. 413 Tipulidae), less often Heteroptera (Anthocoridae), (54045’12”N & 43016’54”E); 2 female, 6.ix.2014, cord. leafhoppers (Cicadellidae), and barkflies (Psocidae). Inorski (54044’15”N & 43008’53”E). Predators: velvet ants Myrmosa atra Panzer, 1801 General distribution: Russia (the European part (Mutillidae), and ichneumon Phygadeuon rugulosus from the centre till Crimea and northern Caucasus, Gravenhorst, 1829 (Ichneumonidae). Urals, Tomsk region, Altai, eastern Siberia, Amur region, and Primorsky Krai), Europe, northern Africa, Turkey, borealis (Zetterstedt, 1838) Kazakhstan, Uzbekistan, Tajikistan, Iran, and Mongolia Literature: Mokrousov et al. 2013; Ruchin & (Antropov 2017b). Antropov 2014. Biology: Nests in sandy or clay soil, often in Material: RM: 2 female, 27.vii.2014, 27.v.2016, groups. Prey: various small dipterans (Anisopodidae MSNR, quar. 360 (54046’13”N & 43013’31”E); 1 female,

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13.viii.2015, quar. 249 (54049’39”N & 43032’08”E); 1 Ectemnius cephalotes (Olivier, 1792) female, 7.vi.2015, quar. 399 (54046’40”N & 43027’27”E); Literature: Mokrousov et al. 2009, 2013; Feoktistov 1 male, 13.vii.2015, quar. 434 (54045’04”N & 43019’02”E); 2011; Ruchin & Antropov 2014. 1 male, 5.vii.2015, quar. 405; 1 female, 28.vii.2015, quar. Material: RM: 1 female, 4.vii.2015, MSNR, quar. 82 (54053’12”N & 43032’46”E); 1 male, 17.viii.2014, quar. 288 (54048’27”N & 43022’03”E); 1 female, 7.ix.2014, 397 (54046’17”N & 43025’44”E); 1 male, 5.vii.2015, quar. quar. 420 (54045’29”N & 43024’19”E); 1 female, 403 (54044’40”N & 43006’57”E); 1 female, 23.viii.2016, 24.viii.2017, quar. 368 (54046’37”N & 43021’45”E); 2 quar. 85 (54053’29”N & 43035’00”E); 1 female, female, 5.vii.2014, 16.viii.2015, Pushta (54042’50”N & 18.vii.2016, quar. 383 (54045’36”N & 43011’48”E); 1 43013’32”E). male, 15.vi.2016, quar. 409; 1 male, 4.vii.2017, quar. General distribution: Russia (the European part from 357 (54046’07”N & 43010’56”E); 1 female, 8.vi.2014, the northwest till Crimea and northern Caucasus, Urals, cord. Inorski (54044’15”N & 43008’53”E); 1 female, Altai, and Primorsky Krai), Europe, northern Africa, 19.vii.2016, cord. Podrubnyi (54047’51”N & 43008’48”E); Turkey, Syria, Kazakhstan, and North America (Antropov NNR: 1 ex., 29.v.2015, Sarov (54055’08” N 43017’40”E), 2017b). coll. P.V. Cherenkov (Image 3). Biology: Gnaw nests in rotten wood, but sometimes General distribution: Russia (the European part in dense wood. They often settle in groups in which from the north till Crimea and northern Caucasus, different females can use a common entrance. Prey: Urals, Altai, eastern Siberia, Amur region, Kamchatka, various flies (Calliphoridae, Muscidae, Syrphidae, Primorsky Krai, Sakhalin, and Kuril Islands), Europe, Tabanidae) (Giovanni et al. 2017). Turkey, Kazakhstan, Mongolia, Korea, Japan (Hokkaido), and North America (Antropov 2017b). Ectemnius continuus (Fabricius, 1804) Biology: Nests in rotten wood. Prey: various small Literature: Redikortsev 1938 (as Crabro vagus L.); flies. Plavilshchikov 1964 (as Crabro vagus L.); Ruchin et al. 2009; Mokrousov et al. 2013; Ruchin & Antropov 2014. Ectemnius cavifrons (Thomson, 1870) Material: RM: 1 female, 1 male, 28.vii.2015, Literature: Plavilshchikov 1964 (as Crabro plumifrons MSNR, quar. 79 (54053’11”N & 43029’18”E); 2 female, Thms.); Mokrousov et al. 2009, 2013; Ruchin & Antropov 13.viii.2015, quar. 249 (54049’39”N & 43032’08”E); 2014. 1 female, 1.vii.2016, quar. 379 (54045’17”N & Material: RM: 1 male, 24.vii.2016, MSNR, quar. 398 43007’51”E); 1 male, 15.vii.2017, quar. 324 (54046’28”N 0 0 (54 46’22”N & 43 25’48”E); 1 male, 19.vii.2016, cord. & 43005’40”E); 1 female, 6.viii.2017, quar. 397 0 0 Podrubnyi (54 47’51”N & 43 08’48”E). (54046’17”N & 43025’44”E); 1 female, 13.viii.2016, quar. General distribution: Russia (the European part from 446 (54043’13”N & 43013’23”E); 1 female, 17.viii.2014, the northwest till Crimea and northern Caucasus, Urals, cord. Polyanski (54046’36”N & 43028’47”E); 1 male, Siberia, Amur region, Khabarovsk Krai and Primorsky 12.ix.2017, cord. Plotomoika (54053’36”N & 43009’41”E); Krai, Sakhalin, and Kuril Islands), Europe, northern 1 female, 10.viii.2014, Temnikov Distr., Tretyakovo Africa, Kazakhstan, Tajikistan, Mongolia, Korea, Japan (54031’44”N & 43013’18”E); 1 female, 24.vii.2016, (Hokkaido, Honshu, Shikoku, Kyushu, and Ryukyu), and Polyanki (54044’27”N & 43029’04”E); 1 male, 23.vii.2016, southeastern Asia (Antropov 2017b). Elniki Distr., Cherlyaj (54045’53”N & 43040’58”E); NNR: 1 Biology: Gnaws nests in rotten wood. Prey: various female, 1.viii.2017, Pervomaisk Distr., 6km southwest of dipterans (mainly Episyrphus balteatus (De Geer, Lesozavod (54054’32”N & 43038’11”E). 1776) (Syrphidae), Calliphoridae, Sarcophagidae, General distribution: Russia (the European part Stratiomyiidae, and Tabanidae) and leafhoppers from the northwest till Crimea and northern Caucasus, Iassus lanio Linnaeus, 1761 (Cicadellidae) (Giovanni Urals, Siberia, Amur region, Kamchatka, Khabarovsk et al. 2017). Predators: chalcid wasps Pteromalus Krai and Primorsky Krai, Sakhalin, and Kuril Islands), (Pteromalidae), ichneumons Perithous albicinctus Europe, northern Africa, Israel, Jordan, Turkey, Georgia, (Gravenhorst, 1829) (Ichneumonidae), emerald wasps Azerbaijan, Syria, Kazakhstan, central Asia, Iraq, Iran, Chrysis fulgida Linnaeus, 1761 (Chrysididae), and Mongolia, southwestern and northern China, Korea, parasitic flies Eustalomyia festiva (Zetterstedt, 1845) and Japan (Hokkaido, Honshu, Shikoku, and Kyushu). and E. hilaris (Fallen, 1823) (Anthomyiidae). As a result of accidental importation, the species acclimatized in North America, Cuba, Guatemala, and

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Venezuela (Antropov 2017b). northeastern China, Korea, and Japan (Hokkaido and Biology: The species gnaws nests in rotten wood Honshu) (Antropov 2017b). or in dry branches with a soft core. Prey: various flies Biology: Gnaws nests mostly in rotten wood, but (Anthomyiidae, Calliphoridae, Muscidae, Rhagionidae, often in dense wood, as well as in clay walls. Prey: large Sciomyzidae, Stratiomyidae, Syrphidae, Tabanidae, flies Syrphidae and Asilidae. Tachinidae, and Therevidae) (Giovanni et al. 2017). Predators: ichneumons Caenocryptus inflatus(Thomson, Ectemnius guttatus (Vander Linden, 1829) 1884) and Hoplocryptus confector (Gravenhorst, 1829) Literature: Redikortsev 1938 (as Crabro spinicollis H. (Ichneumonidae) and parasitic fliesMacronychia aurata Sch.); Plavilshchikov 1964 (as Crabro spinicollis H.-Sch.). (Coquillett, 1902) (Sarcophagidae). Material: RM: 1 female, 17.vi.2016, MSNR, quar. 330 (54046’44”N & 43011’09”E); 1 female, 18.vii.2016, quar. Ectemnius dives (Lepeletier de Saint Fargeau & Brullé, 303 (54047’17”N & 43012’32”E); 2 male, 4.vi.2016, quar. 1835) 383 (54045’36”N & 43011’48”E). Literature: Plavilshchikov 1964 (as Crabro dives General distribution: Russia (the European part Lep.); Mokrousov et al. 2013. from the northwest till Crimea and northern Caucasus, Material: RM: 1 male, 12.vi.2013, MSNR, cord. Urals, Altai, Irkutsk region, and Transbaikalia), Europe, Podrubnyi (54047’51”N & 43008’48”E). and Kazakhstan (Antropov 2017b). General distribution: Russia (the European part from Biology: Gnaw nests in rotten wood or in shoots the northwest till Crimea and northern Caucasus, Urals, with a soft core. Prey – various dipterans (Chloropidae, Siberia, Amur region, Khabarovsk Krai and Primorsky Muscidae, and Syrphidae). Krai, Sakhalin, and Kuril Islands), Europe, northern Africa, Remark: The species is confirmed in MSNR. Israel, Turkey, Georgia, Kazakhstan, southwestern China, Korea, Japan (Hokkaido and Honshu), and North Ectemnius lapidarius (Panzer, 1804) America (Antropov 2017b). Literature: Redikortsev 1938 (Crabro chrysostomus Biology: The species gnaws nests in rotten Lep.); Plavilshchikov 1964 (Crabro chrysostomus Lep.); wood. Prey: various flies (Syrphidae and Tachinidae). Ruchin et al. 2009; Mokrousov et al. 2013; Ruchin & Predators: emerald wasps Chrysis grohmanni Dahlbom, Antropov 2014. 1854 [Chrysididae] and ichneumons Perithous mediator Material: RM: 1 female, 28.vii.2015, MSNR, quar. 82 (Fabricius, 1804) (Ichneumonidae). (54053’12”N & 43032’46”E); 1 male, 11.vii.2015, quar. Remark: The species is confirmed in MSNR. 142 (54051’12”N & 43009’21”E); 1 male, 27.vii.2014, quar. 360 (54046’13”N & 43013’31”E); 1 male, 21.vi.2015, Ectemnius fossorius (Linnaeus, 1758) quar. 384 (54045’33”N & 43012’44”E); 2 female, Literature: Mokrousov et al. 2013; Ruchin & 23.viii.2015, quar. 387 (54045’43”N & 43015’46”E); 1 Antropov 2014. female, 7.vi.2015, quar. 399; 1 male, 21.v.2015, quar. 408 Material: RM: 1 female, 28.vii.2015, MSNR, quar. 79 (54044’57”N & 43011’47”E); 1 female, 5.vi.2015, quar. (54053’11”N & 43029’18”E); 1 female, 28.vii.2015, quar. 448 (54042’37”N & 43012’29”E); 1 male, 11.vii.2015, 82 (54053’12”N & 43032’46”E); 1 female, 9.viii.2014, quar., 142 (54051’12”N & 43009’21”E); 2 male, 1 female, quar. 389 (54045’50”N & 43017’50”E); 1 female, 1 male, 25.v.2014, 19.vii.2016, cord. Podrubnyi (54047’51”N 17.vi.2014, quar. 408 (54044’57”N & 43011’47”E); 2 & 43008’48”E); 1 female, 6.vii.2013, Temnikov Distr., male, 18.vii.2016, quar. 383 (54045’36”N & 43011’48”E); Lavrentevo (54029’44”N & 43002’41”E); 1 female, 2 female, 15.viii.2014, 16.viii.2014, cord. Taratinski 19.vii.2015, Elniki Distr., Malye Mordovskie Poshaty (54044’43”N & 43005’14”E); 1 male, 11.vii.2015, cord. (54041’00”N & 43043’50”E). Podrubnyi (54047’51”N & 43008’48”E); 1 female, General distribution: Russia (the European part from 7.viii.2017, cord. Pavlovski (54045’15”N & 43024’03”E), the northwest till Crimea and northern Caucasus, Urals, coll. G.B. Semishin; 1 male, 23.viii.2017, cord. Inorski Siberia, Amur region, Khabarovsk Krai and Primorsky (54044’15”N & 43008’53”E), coll. G.B. Semishin. Krai, Sakhalin, and Kuril Islands), Europe, Turkey, General distribution: Russia (the European part from Kazakhstan, Kyrgyzstan, Iran, Mongolia, northeastern the northwest till Crimea and northern Caucasus, Urals, China, Korea, Japan (Hokkaido and Honshu), Africa, and Siberia, Amur region, Khabarovsk Krai and Primorsky North America (Antropov 2017b). Krai, Sakhalin, and Kuril Islands), Europe, northern Africa, Biology: Gnaws nests in rotten wood. Prey: Turkey, Kazakhstan, Uzbekistan, Kyrgyzstan, Mongolia, various dipterans (Calliphoridae, Empididae, Muscidae,

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Stratiomyidae, Sphaerophoria scripta (Linnaeus, 1758) (), Anthomyiidae, Cyrtidae. Dolichopodidae, (Syrphidae), and Therevidae). Predators: chalcid Muscidae, Stratiomyidae, Syrphidae, Tachinidae, and wasps Pteromalus and Habritys brevicornis (Ratzeburg, Trypetidae) and sometimes mayflies (Ephemeroptera). 1844) [Pteromalidae], ichneumons Perithous mediator Predators: emerald wasps Trichrysis cyanea (Linnaeus, (Fabricius, 1804) (Ichneumonidae), and parasitic flies 1758) (Chrysididae), velvet ants Cystomutilla ruficeps Eustalomyia hilaris (Fallen, 1823) (Anthomyiidae). (Smith, 1855) and Myrmosa brunnipes (Lepeletier de Saint Fargeau, 1845) (Mutillidae), chalcid wasps Ectemnius lituratus (Panzer, 1805) Torymus cupreus (Spinola 1808) () and Literature: Plavilshchikov 1964 (Crabro lituratus Eurytoma nodularis Boheman, 1836 (), Panz.); Mokrousov et al. 2013. as well as ichneumons Endasys sugiharai (Uchida, Material: RM: 1 female, 1 male, 13.vii.2015, MSNR, 1936), Hoplocryptus, Kaltenbachia, and Scambus quar. 434 (54045’04”N & 43019’02”E); 1 female, 2 male, (Ichneumonidae). 15.viii.2014, cord. Taratinski (54044’43”N & 43005’14”E); Remark: The species is confirmed in MSNR. 1 male, 11.vii.2015, cord. Podrubnyi (54047’51”N & 43008’48”E). Ectemnius ruficornis (Zetterstedt, 1838) General distribution: Russia (the European part Literature: Redikortsev 1938 (as Crabro plumifrons from the centre till Crimea and northern Caucasus and Thom.); Mokrousov et al. 2013. Urals), Europe, northern Africa, Turkey, Kazakhstan, and Material: RM: 1 female, 5.vi.2015, MSNR, quar. 434 Iran (Antropov 2017b). (54045’04”N & 43019’02”E); 1 male, 21.vi.2015, quar. 360 Biology: The species gnaws nests in rotten wood. (54046’13”N & 43013’31”E); 1 male, 5.vi.2015, quar. 448 Prey: dipterans (Anthomyiidae and Sarcophapidae) (54042’37”N & 43012’29”E); 1 male, 18.vii.2016, quar. (Giovanni et al. 2017). 383 (54045’36”N & 43011’48”E); 1 male, 11.vi.2016, quar. 389 (54045’50”N & 43017’50”E); 1 male, 15.vi.2016, Ectemnius nigritarsus (Herrich-Schäffer, 1841) quar. 429 (54044’31”N & 43014’59”E); 1 female, 2 male, Literature: Mokrousov et al. 2013. 11.vii.2015, 19.vii.2016, cord. Podrubnyi (54047’51”N & General distribution: Russia (the European part from 43008’48”E). the centre till Crimea, Urals, Kemerovo region, Altai, General distribution: Russia (the European part Amur region, and Primorsky Krai), Europe, Georgia, from the northwest till Crimea and northern Caucasus, Turkey, India, Sri Lanka, Korea, Japan (Hokkaido and Urals, Siberia, Amur region, Kamchatka, Khabarovsk Krai Honshu), southeastern Asia, and Australia (Antropov and Primorsky Krai, Sakhalin, and Kuril Islands), Europe, 2017b). Turkey, Kazakhstan, Mongolia, Korea, Japan (Hokkaido, Biology: Gnaws nests in rotten wood. Prey: various Honshu, and Ryukyu), southeastern Asia, and North dipterans (Anthomyiidae, Muscidae, Syrphidae, America (Antropov 2017b). Tachinidae, and Therevidae). Biology: Nests in rotten wood. Prey: various dipterans (Anthomyiidae, Dolichopodidae, Muscidae, Ectemnius rubicola (Dufour et Perris, 1840) and Syrphidae (Episyrphus balteatus (De Geer, 1776) Literature: Redikortsev 1938 (as Crabro larvatus and Syrphus torvus Osten Sacken, 1875)). Predators: Wesm.); Plavilshchikov 1964 (as Crabro larvatus Venn.); emerald wasps Pseudomalus auratus (Linnaeus, Ruchin & Antropov 2016. 1758) (Chrysididae) and ichneumons Perithous scurra Material: RM: 1 female, 12.vi.2016, MSNR, quar. 430 japonicus Uchida, 1928 (Ichneumonidae). (54044’34”N & 43016’00”E). Remark: The species is confirmed in MSNR. General distribution: Russia (the European part from the northwest till Crimea and northern Caucasus, Ectemnius spinipes (A. Morawitz, 1866) Urals, Altai, Kemerovo, Irkutsk and Amur regions, Literature: Ruchin et al. 2009; Mokrousov et al. Primorsky Krai, Sakhalin, and Kuril Islands), Europe, 2013; Ruchin & Antropov 2014. Turkey, Kazakhstan, Iran, and Japan (Hokkaido, Honshu, Material: RM: 1 male, 13.vi.2016, MSNR, quar. 421 Shikoku, and Kyushu) (Antropov 2017b). (54043’52’’N, 43007’06”E); 1 male, 18.vii.2016, quar. 383 Biology: The species gnaws nests in shoots with (54045’36”N & 43011’48”E); 2 male, 13.vi.2016, cord. soft core of both dry and live plants, hollow stems of Inorski (54044’15”N & 43008’53”E). herbaceous plants, or rotten wood. Prey: various General distribution: Russia (the European part from dipterans ( gibbosus (Linnaeus, 1758) the centre till Crimea and northern Caucasus, Urals,

13230 Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov

Siberia, Amur region, Khabarovsk Krai and Primorsky Krai, Antropov 2014. and Sakhalin), Europe, Turkey, Azerbaijan, Kazakhstan, Material: RM: 1 female, 28.vii.2015, MSNR, quar. Turkmenistan, Mongolia, northeastern China, Korea, 79 (54053’11”N & 43029’18”E); 3 female, 28.vii.2015, and Japan (Hokkaido and Honshu) (Antropov 2017b). quar. 82 (54053’12”N & 43032’46”E); 8 female, 1 Biology: Gnaws nests in rotten wood. Prey: various male, 11.vii.2015, 18.vi.2016, 12.ix.2017, quar. 115 small lepidopterans (Drepanidae, Geometridae, (54051’46”N & 43009’26”E); 1 female, 13.viii.2015, quar. Noctuidae, Pyralidae, Tortricidae, Uraniidae, and, 249 (54049’39”N & 43032’08”E); 3 female, 13.viii.2015, sometimes, Lycaenidae). Predators: ichneumons quar. 274; 6 female, 20.vii.2014, 11.vii.2015, quar. 301 Exeristes (Ichneumonidae). (54047’19”N & 43010’36”E); 1 female, 4.vii.2015, quar. 368 (54046’37”N & 43021’45”E); 1 female, 23.viii.2015, ** Lestica alata (Panzer, 1797) quar. 387 (54045’43”N & 43015’46”E); 1 female, Material: RM: 1 female, 15.viii.2015, Temnikov 9.viii.2014, quar. 389 (54045’50”N & 43017’50”E); 1 Distr., Purdoshki (54040’04”N & 43032’33”E). female, 9.viii.2014, quar. 413 (54045’12”N & 43016’54”E); General distribution: Russia (the European part from 2 female, 1 male, 20.vii.2014, 4.vii.2017, quar. 276 the northwest till Crimea and northern Caucasus, Urals, (54047’45”N & 43010’28”E); 1 male, 27.vii.2014, quar. Siberia, Amur region, and Khabarovsk Krai and Primorsky 360 (54046’13”N & 43013’31”E); 3 male, 27.vii.2014, Krai), Europe, northern Africa, Turkey, Kazakhstan, 21.vi.2015, 14.vi.2016, quar. 384 (54045’33”N & Kyrgyzstan, Mongolia, north of India, northern China, 43012’44”E); 2 male, 17.vi.2014, 11.vii.2017, quar. 408 Korea, and Japan (Hokkaido) (Antropov 2017b). (54044’57”N & 43011’47”E); 1 male, 13.vii.2015, quar. Biology: Nests in the soil. Prey: various small 434 (54045’04”N & 43019’02”E); 1 male, 5.vii.2015, lepidopterans (Agriphila geniculea (Haworth, 1811), quar. 405; 1 male, 16.vii.2015, quar. 442 (54043’56”N & Agriphila tristellus (Denis et Schiffermüller, 1775), 43015’02”E); 1 male, 23.vi.2013, quar. 431 (54044’31”N Chrysoteuchia culmella Linnaeus, 1758), Crambus & 43017’14”E); 1 female, 1 male, 15.vi.2016, quar. 429 pascuellus Linnaeus, 1758) (Crambidae), Geometridae, (54044’31”N & 43014’59”E); 2 male, 18.vi.2016, quar. Noctuidae, Pyralidae, and Tortricidae). 61; 1 female, 1 male, 1.vii.2016, quar. 329 (54046’36”N & 43010’39”E); 1 male, 7.vii.2016, quar. 361; 3 male, Lestica camelus (Eversmann, 1849) 1.vii.2016, quar. 379 (54045’17”N & 43007’51”E); Literature: Mokrousov et al. 2013. 1 female, 18.vii.2016, quar. 331 (54046’43”N & Material: RM: 1 female, 13.viii.2015, MSNR, quar. 400 43012’37”E); 5 female, 3 male, 18.vii.2016, quar. (54047’18”N & 43029’06”E); 1 female, 9.viii.2014, quar. 358 (54046’05”N & 43011’44”E); 2 female, 2 male, 413 (54045’12”N & 43016’54”E); 1 female, 18.vii.2016, 18.vii.2016, quar. 383 (54045’36”N & 43011’48”E); 1 quar. 383 (54045’36”N & 43011’48”E); 1 male, 1.vii.2016, female, 2 male, 4.vii.2017, quar. 357 (54046’07”N & quar. 406 (54045’07”N & 43009’54”E); 2 female, 43010’56”E); 2 male, 13.vii.2017, quar. 319 (54048’09”N 1.viii.2017, quar. 111 (54052’59”N & 43033’01”E); 1 male, & 43028’25”E); 1 female, 1 male, 18.vii.2017, quar. 2.viii.2016, cord. Inorski (54044’15”N & 43008’53”E); 380 (54045’06”N & 43008’34”E); 1 female, 1.viii.2017, 1 female, 22.vii.2017, Elniki Distr., Malye Mordovskie quar. 111; 3 male, 9.vi.2013, 11.vii.2015, 4.vii.2017, Poshaty (54041’00”N & 43043’50”E); NNR: 1 male, cord. Podrubnyi (54047’51”N & 43008’48”E); 2 male, 1.viii.2017, 6km southwest of Lesozavod (54054’32”N & 28.vi.2015, 17.viii.2013, cord. Inorski (54044’15”N & 43038’11”E). 43008’53”E). General distribution: Russia (the centre and the General distribution: Russia (the European part east of the European part, Urals, Siberia, Amur region, from the northwest till Crimea and northern Caucasus, Khabarovsk Krai and Primorsky Krai, and Sakhalin), Urals, Siberia, and Primorsky Krai), Europe, northern Turkey, Kazakhstan, Mongolia, southwestern and Africa, Israel, Lebanon, Turkey, Georgia, Kazakhstan, northeastern China, Korea, and Japan (Hokkaido and Uzbekistan, Kirghizia, Turkmenistan, Syria, Iraq, and Iran Honshu) (Antropov 2017b). (Antropov 2017b). Biology: Nests in the abandoned tunnels of Biology: Nests in the abandoned tunnels of xylophagous insects in dry wood. Prey: various leafroller xylophagous insects. Prey: various lepidopterans moths (Tortricidae). (Crambidae, Noctuidae, Sesiidae, and Sterrhidae) and sometimes dipterans (Empididae, Muscidae, and Lestica clypeata (Schreber, 1759) Syrphidae). Predators: emerald wasps Chrysis fasciata Literature: Mokrousov et al. 2013; Ruchin & Olivier, 1790 (Chrysididae) and digger wasps Palarus

Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 13231 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov variegatus (Fabricius, 1781) (Crabronidae). 420 (54045’29”N & 43024’19”E). General distribution: Russia (the European part ** Lestica subterranea (Fabricius, 1775) from the centre to northern Caucasus and Urals), Material: RM: 1 male, 18.vi.2016, MSNR, quar. 197 Europe, northern Africa, Israel, Jordan, Turkmenistan, (54050’05”N & 43009’09”E). Syria, Kazakhstan, central Asia, Iran, and Afghanistan General distribution: Russia (the European part (Antropov 2017b). from the northwest till Crimea and northern Caucasus, Biology: Nests in dense soil. Prey: small Urals, and Altai), Europe, northern Africa, Turkey, hymenopterans (, Formicidae, Kazakhstan, Kyrgyzstan, Tajikistan, Iran, and northern Ichneumonidae, and Pteromalidae) and sometimes China (Antropov 2017b). dipterans (Ceratopogonidae and Lycoriidae). Biology: Nests in the soil. Prey: various small Remark: The species is confirmed in MSNR. lepidopterans (Crambidae, Lythria purpuraria Linnaeus, 1758 (Geometridae), Glyphipterigidae, and Tortrix Tribe Larrini viridana Linnaeus, 1758 (Tortricidae), rarely Zygaenidae). * Tachysphex ferrugineus Pulawski, 1971 Material: RM: 1 female, 2.vii.2017, MSNR, cord. 0 0 Lindenius albilabris (Fabricius, 1793) Inorski (54 44’15”N & 43 08’53”E). Literature: Redikortsev 1938 (Crabro albilabris F.); General distribution: Russia (the European part Plavilshchikov 1964 (Crabro albilabris F.); Ruchin et al. from the centre to northern Caucasus), eastern Europe, 2009; Ruchin & Antropov 2016. Turkey, and Kazakhstan (Antropov 2017b). General distribution: Russia (the European part from Tachysphex fulvitarsis (Costa, 1867) the northwest till Crimea and northern Caucasus, Urals, Literature: Mokrousov et al. 2013. Siberia, Amur region, and Khabarovsk Krai and Primorsky Material: RM: 1 male, 4.viii.2015, MSNR, quar. 86 Krai), Europe, northern Africa, Israel, Turkey, Syria, (54053’38”N & 43035’59”E). Kazakhstan, Uzbekistan, Kyrgyzstan, Turkmenistan, General distribution: Russia (the European part Mongolia, and Korea (Antropov 2017b). from the centre till Crimea and northern Caucasus, Biology: Nests in dense gravel or sandy soil. Prey: Urals, Siberia, Amur region, and Primorsky Krai), heteropterans Miris (Miridae) or small dipterans Europe, northern Africa, Israel, Jordan, Turkey, Syria, (Chloropidae, Dolichopodidae, and Muscidae). Kazakhstan, Uzbekistan, Turkmenistan, and Tajikistan Predators: emerald wasps Hedychridium coriaceum (Antropov 2017b). (Dahlbom, 1854) (Chrysididae) and velvet ants Myrmosa Biology: Nests in soil. Prey: hoppers Metrioptera atra Panzer, 1801 (Mutillidae). bicolor (Philippi, 1830), Platycleis, and Roeseliana roeselii (Hagenbach, 1822) (Tettigoniidae). Lindenius panzeri (Vander Linden, 1829) Remark: The species is confirmed in MSNR. Literature: Redikortsev 1938 (Crabro panzeri Rind.); Plavilshchikov 1964 (Crabro panzeri Lind.); Ruchin et al. Tachysphex helveticus Kohl, 1885 2009; Mokrousov et al. 2013. Literature: Mokrousov et al. 2013. General distribution: Russia (the European part General distribution: Russia (the European part from from the centre to northern Caucasus, Urals, Siberia, the centre to northern Caucasus, Urals, Altai, eastern Amur region, and Khabarovsk Krai and Primorsky Krai), Siberia, and Primorsky Krai), Europe, northern Africa, Europe, northern Africa, Turkey, Kazakhstan, Uzbekistan, Jordan, Turkey, Kazakhstan, Tajikistan, and Mongolia Kyrgyzstan, Turkmenistan, Iran, Afghanistan, Mongolia, (Antropov 2017b). and northeastern China (Antropov 2017b). Biology: Nests in soil. Prey: grasshoppers Calliptamus Biology: Nests in sandy or clay soil. Prey: small italicus (Linnaeus, 1758) (Acrididae). Predators: velvet dipterans (Chloropidae, Ephydridae, and Trypetidae). ants Smicromyrme rufipes (Fabricius, 1787) (Mutillidae). Predators: velvet ants Myrmosa atra Panzer, 1801 (Mutillidae). Tachysphex obscuripennis (Schenck, 1857) Literature: Mokrousov et al. 2013. Lindenius pygmaeus (Rossi, 1794) Material: RM: 1 male, 2.vii.2017, MSNR, cord. Inorski Literature: Mokrousov et al. 2013. (54044’15”N & 43008’53”E). Material: RM: 1 female, 25.vii.2013, MSNR, quar. General distribution: Russia (the European part from

13232 Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov the centre till northern Caucasus and Altai), Europe, and Melanoplus (Acrididae). Morocco, Lebanon, and Turkey (Antropov 2017b). Biology: Nests in soil. Prey: cockroaches Ectobius Tachytes panzeri (Dufour, 1841) lapponicus (Linnaeus, 1758) and E. pallidus (Olivier, 1789) Literature: Ruchin et al. 2009; Mokrousov et al. 2013. (Ectobiidae). Predators: emerald wasps Chrysis bicolor Material: RM: 1 female, 19.vii.2015, Elniki Distr., 0 0 Lepeletier de Saint Fargeau, 1806 and Hedychridium Malye Mordovskie Poshaty (54 41’00”N & 43 43’50”E). ardens (Coquebert, 1801) (Chrysididae). General distribution: Russia (the European part Remark: The species is confirmed in MSNR. from the centre till Crimea and northern Caucasus, Urals, Novosibirsk region, Altai, and Irkutsk region), Tachysphex pompiliformis (Panzer, 1805) south of Europe, northern Africa, Israel, Turkey, Kazakhstan, Uzbekistan, Turkmenistan, Tajikistan, and Literature: Mokrousov et al. 2013; Ruchin & Iran (Antropov 2017b). Antropov 2014. Biology: Nests in sandy soil. Prey: nymphs of Material: RM: 1 male, 19.vii.2015, Elniki Distr., Malye grasshoppers Oedipoda spp. or Stenobothrus sp. Mordovskie Poshaty (54041’00”N & 43043’50”E). (Acrididae). Predators: brood parasites Brachystegus General distribution: Russia (the European part scalaris (Illiger, 1807) and female digger wasps Palarus from the north till Crimea and northern Caucasus, variegatus (Fabricius, 1781) (Crabronidae). Urals, Siberia, Amur and Magadan regions, Kamchatka, Remark: The species is confirmed in the territories Khabarovsk Krai and Primorsky Krai, and Sakhalin), adjacent to MSNR. Europe, northern Africa, Turkey, Kazakhstan, Uzbekistan, Turkmenistan, Tajikistan, Iran, Pakistan, India, Mongolia, Tribe Miscophini northern China, Korea, and North America (Antropov Miscophus ater Lepeletier, 1845 2017b). Literature: Mokrousov et al. 2013. Biology: Nests in soil. Prey: nymphs of grasshoppers General distribution: Russia (the centre and south Acrotylus, Aulocara, Calliptamus coelesyriensis Giglio- of the European part, Altai, Irkutsk region, Amur region, Tos, 1893, Camnula, Chorthippus, Dericorys tibialis and Primorsky Krai), Europe, northern Africa, Turkey, (Pallas, 1773), Dociostaurus kraussi (Ingenizkij, 1897), Azerbaijan, Kazakhstan, and Iran (Antropov 2017b). Gomphocerippus rufus (Linnaeus, 1758), Gomphocerus, Biology: Nests in soil. Prey: juvenile spiders Salticidae Melanoplus, Oedaleonotus, Omocestus, and and Theridiidae. Stenobothrus (Acrididae), sometimes nymphs of hoppers (Tettigoniidae). Predators: parasitic flies Apodacra, Miscophus bicolor Jurine, 1807 Taxigramma, and Xeromyіa (Sarcophagidae), emerald Literature: Mokrousov et al. 2013. wasps Hedychridium adventicium Zimmermann, 1962, General distribution: Russia (the European part from H. ardens (Coquebert, 1801), H. roseum (Rossi, 1790), the centre to northern Caucasus, Urals, Altai, eastern Siberia, Amur region, Khabarovsk Krai and Primorsky Chrysis bicolor Lepeletier de Saint Fargeau, 1806, C. Krai, and Sakhalin), Europe, northern Africa, Turkey, illigeri Wesmael, 1839, and C. succincta Linnaeus, 1767 Syria, Abkhazia, Kazakhstan, Iran, Mongolia, and Korea (Chrysididae), and velvet ants Smicromyrme sicanus (De (Antropov 2017b). Stefani, 1887) (Mutillidae). Biology: Nests in sandy soil. Prey: juvenile spiders [Theridiidae]. Predators: emerald wasps Chrysis cortii * Tachysphex psammobius (Kohl, 1880) Linsenmaier, 1951, C. lanceolata Linsenmaier, 1959, C. Material: RM: 1 male, 30.v.2015, Temnikov Distr., leachii Shuckard, 1836, and Hedychridium elegantulum 0 0 Alkaevo (54 35’49”N & 43 22’35”E). du Buysson, 1887 (Chrysididae) and velvet ants General distribution: Russia (the European part Smicromyrme sicanus (De Stefani, 1887) (Mutillidae). from the centre till Crimea and northern Caucasus, Urals, Tomsk region, Altai, Irkutsk region, Buryatia, and Miscophus concolor Dahlbom, 1844 Primorsky territory), south of Europe, northern Africa, Literature: Mokrousov et al. 2013. Israel, Turkey, Kazakhstan, China, and North America General distribution: Russia (the centre and the east (Antropov 2017b). of the European part) and Europe (Antropov 2017b). Biology: Nests in sandy soil. Prey: nymphs of Biology: Nests in sandy soil. Prey: juvenile grasshoppers Circotettix maculatus S.H. Scudder, 1880 spiders Salticus (Salticidae) and Stemonyphantes

Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 13233 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov lineatus (Linnaeus, 1758) (Linyphiidae). Predators: Europe, northern Africa, Turkey, Syria, Iran, Kazakhstan, emerald wasps Hedychridium zelleri (Dahlbom, 1845) Uzbekistan, Turkmenistan, Morgolia, Korea, and North (Chrysididae). America (Antropov 2017b). Biology: Nests in loose sandy soil, sometimes in Miscophus johni Mokrousov, 2004 small groups. Prey: various dipterans Calythea nigricans Literature: Mokrousov et al. 2013. (Robineau-Desvoidy, 1830) (Anthomyiidae), Pollenia General distribution: Russia (the centre of the rudis (Fabricius, 1794) (Calliphoridae), Fannia incisurata European part) and southeast of Europe (Antropov (Zetterstedt, 1838) (Fannidae), Hydrotaea armipes 2017). (Fallén, 1825), Myospila meditabunda (Fabricius, 1781) (Muscidae), Leucostoma aterrimum (Villers, 1789), ? Nitela spinolae Latreille, 1809 Strongygaster triangulifer (Loew, 1863) (Tachinidae), Literature: Plavilshchikov 1964 (as Nitela spinolai as well as Chloropidae, Dolichopodidae, Lonchaeidae, Dahlb.). Milichiidae, Pipunculidae, Platypezidae, Rhagionidae, General distribution: Russia (the European part Sarcophagidae, Stratiomyidae, Syrphidae, and from the north to northern Caucasus), Europe, northern Therevidae. Predators: emerald wasps Hedychridium Africa, Turkey, and Japan (Hokkaido) (Antropov 2017b). ardens (Coquebert, 1801) (Chrysididae), velvet ants Biology: Nests in hollow stalks of cereals, in Smicromyrme lewisi Mickel, 1935 and Smicromyrme abandoned tunnels of xylophagous insects in dry wood, rufipes (Fabricius, 1787) (Mutillidae), chalcid wasps or in galls of gallflies Andricus kollari (Hartig, 1843) Perilampus ruficornis (Fabricius, 1793) (), (Cynipidae). Prey: aphids [Aphididae] or nymphs of and parasitic flies Phrosinella fulvicornis (Coquillett, jumping plant lice (Psyllidae) and dust lice (Psocidae). 1895), Senotainia trilineata (Wulp, 1890), Sphecapata Predators: velvet ants Smicromyrme sicanus (De Stefani, conica Fallen, 1810, and Taxigramma multipunctata 1887) (Mutillidae). (Rondani, 1859) (Sarcophagidae). Remark: Confirmation of the species in Mordovia requires more material. Oxybelus haemorrhoidalis Olivier, 1812 Literature: Mokrousov et al. 2013. Tribe Oxybelini General distribution: Russia (the European part from Oxybelus argentatus Curtis, 1833 the northwest to the south, Urals, Novosibirsk region, Literature: Ruchin & Antropov 2016. Altai, Amur region, and Khabarovsk Krai and Primorsky General distribution: Russia (the European part Krai), Europe, northern Africa, Israel, Turkey, Azerbaijan, from the north to the southeast, Urals, Altai, Irkutsk Kazakhstan, Kyrgyzstan, Uzbekistan, Turkmenistan, region, Transbaikalia, and Primorsky Krai), Europe, and Syria, Iran, Mongolia, northern China, Korea, and Japan Mongolia (Antropov 2017b). (Honshu) (Antropov 2017b). Biology: Nests in dense sandy soil. Prey (which is Biology: Nests in sandy soil. Prey: flies Musca delivered to the nest punctured by sting): dipterans, domesticaLinnaeus, 1758, Stomoxys calcitrans (Linnaeus, mostly Acrosathe annulata (Fabricius, 1805), Thereva 1758) (Muscidae), Sarcophila latifrons (Fallen, 1817), fulva (Meigen, 1804), and Th. nobilitata (Fabricius, and Metopia staegeri (Rondani, 1859) (Sarcophagidae). 1775) (Therevidae). Predators: parasitic flies Metopia Predators: parasitic flies Metopia staegeri (Rondani, argyrocephala (Meigen, 1824) and M. campestris (Fallen, 1859) (Sarcophagidae) and digger wasps Palarus 1810) (Sarcophagidae), velvet ants Smicromyrme rufipes variegatus (Fabricius, 1781) (Crabronidae). (Fabricius, 1787) (Mutillidae), and female digger wasps Remark: Confirmation of the species in Mordovia Palarus variegatus (Fabricius, 1781) (Crabronidae). requires more material.

Oxybelus bipunctatus Olivier, 1812 * Oxybelus mandibularis Dahlbom, 1845 Literature: Mokrousov et al. 2013. Material: RM: 1 male, 1.vii.2016, MSNR, quar. 379 Material: RM: 2 male, 12.vi.2013, 1.vii.2016, MSNR, (54045’17”N & 43007’51”E); 1 male, 4.vii.2017, quar. 357 cord. Podrubnyi (54047’51”N & 43008’48”E); 1 male, (54046’07”N & 43010’56”E). 24.vi.2014, Pushta (54042’50”N & 43013’32”E). General distribution: Russia (the European part General distribution: Russia (the European part from from the centre to the southeast, Urals, Tomsk region, the north to the southeast, Urals, Altai, Irkutsk region, Altai, Irkutsk region, Buryatia, Amur region, Primorsky Buryatia, Amur region, Primorsky Krai, and Sakhalin), territory, and Sakhalin), Europe, northern Africa, Turkey,

13234 Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov

Kazakhstan, Kyrgyzstan, Uzbekistan, Turkmenistan, and northeastern China, and North America (Antropov Mongolia (Antropov 2017b). 2017b). Biology: Nests in sandy soil. Prey: flies from the family Biology: Nests in soil of different density, from Muscidae. Predators: emerald wasps Hedychridium clayey-stony to sandy. Prey: various dipterans coriaceum (Dahlbom, 1854) (Chrysididae). (Anthomyiidae, Bombyliidae, Calliphoridae, Chloropidae, Dolichopodidae, Fanniidae, Muscidae, Platystomatidae, Oxybelus quatuordecimnotatus Jurine, 1807 Rhagionidae, Sarcophagidae, Simuliidae, Stratiomyidae, Literature: Mokrousov et al. 2013. and Syrphidae) and Orthoptera (Steiner, 1979). Material: RM: 1 male, 20.vi.2015, Temnikov Distr., Predators: emerald wasps Hedychridium coriaceum Russkoe Karaevo (54040’55”N, 43013’52”E). (Dahlbom, 1854) (Chrysididae), velvet ants Myrmosa General distribution: Russia (the European part atra Panzer, 1801 and Smicromyrme rufipes (Fabricius, from the northwest till Crimea, Urals, Tomsk region, 1787) (Mutillidae), parasitic flies Anthomyia pluvialis Altai, Krasnoyarsk Krai, Irkutsk region, and Khabarovsk (Linnaeus, 1758), Delia florilega (Zetterstedt, 1845), Krai and Primorsky Krai), Europe, northern Africa, Saudi D. radicum (Linnaeus, 1758), Leucophora cinerea Arabia, Oman, Yemen, Israel, Jordan, Turkey, Syria, Robineau-Desvoidy, 1830 (Anthomyiidae), Fannia Kazakhstan, central Asia, Iran, Afghanistan, Pakistan, incisurata (Zetterstedt, 1838), F. scalaris (Fabricius, Mongolia, and southwestern and northeastern China 1794) (Fanniidae), Megaselia pygmaea (Zetterstedt, (Antropov 2017b). 1848) (Phoridae), Macronychia griseola (Fallen, 1820), Biology: Nests in sandy or rocky-sandy soil. Prey: M. polyodon (Meigen, 1824), Metopia argyrocephala dipterans, mostly Thereva arcuata Loew, 1847 (Meigen, 1824), Miltogramma germari Meigen, 1824, (Therevidae), as well as Th. plebeja (Linnaeus, 1758) and Sphecapata conica Fallen, 1810 (Sarcophagidae). (Therevidae), Bellardia corsicana (Villeneuve, 1911) (Calliphoridae), Tolmerus atricapillus (Fallen, 1814) * Oxybelus variegatus Wesmael, 1852 (Asilidae), Drosophila (Drosophilidae), Phytomyza Material: RM: 1 female, 7.vii.2013, Temnikov Distr., (Agromyzidae), Sapromyza (Lauxaniidae), and Tarkhany. Physiphora (Ulidiidae). General distribution: Russia (the European part from the centre till Crimea, Urals, and Altai), Europe, Turkey, Oxybelus trispinosus (Fabricius, 1787) Kazakhstan, Kyrgyzstan, Uzbekistan, Turkmenistan, Iran, Literature: Redikortsev 1938 (as Oxybelus nigripes and Afghanistan (Antropov 2017b). Ol.); Plavilshchikov 1964 (as Oxybelus nigripes Ol.); Biology: Nests in soil of different density, from clayey- Mokrousov et al. 2013. stony to sandy. Prey: house flies Musca domestica General distribution: Russia (the European part from Linnaeus, 1758 (Muscidae). the centre to northern Caucasus, Urals, Altai, Irkutsk Remark: The species is confirmed in the territories region, Buryatia, and Khabarovsk Krai and Primorsky adjacent to MSNR. Krai), Europe, northern Africa, Turkey, and Kazakhstan (Antropov 2017b). Tribe Palarini Biology: Nests in sandy soil. Prey: dipterans Bellardia Palarus variegatus (Fabricius, 1781) pandia (Walker, 1849), B. viarum (Robineau-Desvoidy, Literature: Mokrousov et al. 2013. 1830), Blondelia nigripes (Fallen, 1810), Pollenia Material: RM: 1 female, 12.vii.2015, Elniki Distr., atramentaria (Meigen, 1826), P. griseotomentosa Novye Shaly (54042’13”N & 43038’25”E). (Jacentkovsky, 1944), P. hungarica Rognes, 1987, Nyctia General distribution: Russia (the European part halterata (Panzer, 1798), Calliphora vicina Robineau- from the centre till Crimea and northern Caucasus, Desvoidy, 1830 (Calliphoridae), and Blaesoxipha Urals, Altai, Krasnoyarsk Krai, and Transbaikalia), south plumicornis (Zetterstedt, 1859) (Sarcophagidae). of Europe, northern Africa, Turkey, Armenia, Georgia, Kazakhstan, Uzbekistan, Turkmenistan, Tajikistan, Oxybelus uniglumis (Linnaeus, 1758) Afghanistan, Pakistan, Mongolia, northern China, and Literature: Mokrousov et al. 2013. India (Antropov 2017b). General distribution: Russia (the European part from Biology: Nests in dense clay or rocky-sandy soil. the northwest till Crimea, Urals, Siberia, Kamchatka, Prey: various bees (Apidae) (Ammobates, Andrena, Apis, Sakhalin, and Kuril Islands), Europe, northern Africa, Camptopeum, Chelostoma, Colletes, Dasypoda, Halictus, Turkey, Kazakhstan, Kyrgyzstan, Tajikistan, Mongolia, Hylaeus, Lasioglossum, Megachile, Melitta, Nomada,

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Nomia, Osmia, Panurgus, Sphecodes, Tetralonia, and A. marmoreus Clerck, 1757, A. quadratus Clerck, 1757, Thyreus), wasps Mutilla and Myrmosa (Mutillidae), Araniella, Larinioides cornutus (Clerck, 1757), Mangora Myzinum and Tiphia (Tiphiidae), Scolia, Trielis and male acalypha (Walckenaer, 1802), Metellina segmentata Campsoscolia (Scoliidae), Eumenes, Odynerus, Polistes, (Clerck, 1757), Neoscona adianta (Walckenaer, 1802), Pterocheilus and Vespula (Vespidae), Ammophila, Singa hamata (Clerck, 1757), S. nitidula C.L. Koch, 1844 Podalonia and Prionyx (Sphecidae), Alysson, Astata, and Zilla (Araneidae), Dictyna arundinacea (Linnaeus, Bembecinus, Cerceris, Crabro, Diodontus, Dryudella, 1758) (Dictynidae), Agyneta rurestris (C.L. Koch, 1836), Ectemnius, Lestica, Liris, Oxybelus, Pemphredon, Bathyphantes, Entelecara erythropus (Westring, 1851), Philanthus, Pison, Tachysphex, Tachytes and male Larra Kaestneria dorsalis (Wider, 1834), Linyphia triangularis (Crabronidae)), flies Formica and Camponotus (Clerck, 1757), Neriene montana (Clerck, 1757), N. (Formicidae), as well as parasitic hymenopterans radiata (Walckenaer, 1841) (Linyphiidae), Salticus Leucospis (Leucospididae), Amblyteles, Barylypa, and scenicus (Clerck, 1757), Synageles venator (Lucas, Ichneumon (Ichneumonidae). Predators: velvet ants 1836) (Salticidae), Tetragnatha [Tetragnathidae], Smicromyrme rufipes (Fabricius, 1787) (Mutillidae) and Enoplognatha ovata (Clerck, 1757), Neottiura parasitic flies Protomiltogramma fasciata (Meigen, bimaculata (Linnaeus, 1767), Parasteatoda simulans 1824) (Sarcophagidae). (Thorell, 1875), P. tepidariorum (C.L. Koch, 1841), Remark: The species is confirmed in the territories Platnickina tincta (Walckenaer, 1802), Simitidion simile adjacent to MSNR. (C.L. Koch, 1836), Theridion pictum (Walckenaer, 1802), Th. sisyphium (Clerck, 1757), Th. varians Hahn, 1833 Tribe Trypoxylini (Theridiidae), and Xysticus (Thomisidae) (Giovanni et Trypoxylon deceptorium Antropov, 1991 al. 2017). Predators: emerald wasps Chrysis fasciata Literature: Mokrousov et al. 2013; Ruchin & Olivier, 1790, C. fulgida Linnaeus, 1761, h. ignita Antropov 2016. (Linnaeus, 1758), C. obtusidens Dufour et Perris, 1840, General distribution: Russia (the European part C. splendidula Rossi, 1790, C. viridula Linnaeus, 1761, from the centre till Crimea and northern Caucasus and Elampus panzeri (Fabricius, 1804), Pseudomalus auratus Altai), Europe, northern Africa, Israel, Lebanon, Turkey, (Linnaeus, 1758), P. pusillus (Fabricius, 1804) and Caucasus, Syria, Iran, Kazakhstan, central Asia, and Trichrysis cyanea (Linnaeus, 1758) (Chrysididae), chalcid Mongolia (Antropov 2017b). wasps Melittobia acasta (Walker, 1839) (), Biology: Nests in hollow stems of plants (Bogusch Eurytoma nodularis Boheman, 1836 and E. verticillata et al. 2015). Prey: juvenile spiders of various families. (Fabricius, 1798) (Eurytomidae), and Monodontomerus Predators: chalcid wasps Melittobia acasta (Walker, vicicellae (Walker, 1847) (Torymidae), ichneumons 1839) (Eulophidae) and ichneumons Aritranis coxator Gasteruption assectator (Linnaeus, 1758), G. jaculator (Tschek, 1871), Ephialtes, and Perithous (Ichneumonidae) (Linnaeus, 1758) and G. opacum (Tournier, 1877) (Olberg 1959). (Gasterutpionidae), Aritranis heliophilus (Tschek,1870), Isadelphus armatus (Gravenhorst, 1829), Nematopodius Trypoxylon figulus (Linnaeus, 1758) formosus Gravenhorst, 1829, Perithous divinator (Rossi, Literature: Mokrousov et al. 2009, 2013. 1790), P. scurra (Panzer, 1804), Poemenia notata Material: RM: 2 female, 27.vii.2014, 14.vi.2015, Holmgren, 1859, Polysphincta, Stenodontus marginellus MSNR, quar. 420 (54045’29”N & 43024’19”E); 1 male, (Gravenhorst, 1829), and Townesia tenuiventris 21.vi.2015, quar. 384 (54045’33”N & 43012’44”E); 1 (Holmgren, 1860) (Ichneumonidae), as well as parasitic male, 18.vi.2017, cord. Drozhdenovski (54044’31”N & flies Eustalomyia hilaris (Fallen, 1823) (Anthomyidae), 43017’24”E). Amobia signata (Meigen 1824), Metopia argyrocephala General distribution: Russia (the European part (Meigen, 1824), and Miltogramma punctatum (Meigen, from the northwest till Crimea and northern Caucasus, 1824) (Sarcophagidae) and skin beetles Megatoma Urals, Siberia, and Primorsky Krai), Europe, northern undata (Linnaeus, 1758) (Dermestidae) (Olberg 1959). Africa, Turkey, Kazakhstan, Kyrgyzstan, Uzbekistan, Tajikistan, Mongolia, northern China, Japan (Hokkaido ? Trypoxylon fronticorne Gussakowskij, 1936 and Honshu), and North America (Antropov 2017b). Literature: Feoktistov 2011. Biology: Nests in hollow stems of plants or in General distribution: Russia (the centre and the abandoned tunnels of xylophagous insects in wood. east of the European part, Krasnoyarsk Krai, and Irkutsk Prey: juvenile spiders Araneus diadematus Clerck, 1757, region), Europe, Belarus, , Azerbaijan, Armenia,

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Turkey, Kazakhstan, India, and Nepal (Antropov 2017b). 1806. Biology: Nests in existing cavities (in hollow stems of Remark: The species is confirmed in MSNR. plants and in the tunnels and nests of other insects in wood and shoots with a soft core). Prey: various small Subfamily Mellininae spiders, mostly juvenile. Mellinus arvensis (Linnaeus, 1758) Remark: Confirmation of the species in Mordovia Literature: Redikortsev 1938; Plavilshchikov 1964; requires more material. Mokrousov et al. 2009, 2013; Ruchin & Antropov 2014. Material: RM: 1 female, 4.ix.2015, MSNR, quar. 420 Trypoxylon medium de Beaumont, 1945 (54045’29”N & 43024’19”E); 1 female, 24.viii.2017, quar. Literature: Mokrousov et al. 2013. 368 (54046’37”N & 43021’45”E); 2 female, 18.ix.2017, General distribution: Russia (the European part from Pushta (54042’50”N & 43013’32”E). the centre to the south and east, Altai, and Krasnoyarsk General distribution: Russia (the European part Krai), Europe, Ukraine, Belarus, Estonia, Turkey, and from the northwest till Crimea and northern Caucasus, Kazakhstan (Antropov 2017b). Urals, Altai, and Irkutsk region), Europe, Turkey, and Biology: Nests in existing cavities (in hollow stems of Kazakhstan (Nemkov 2017d). plants and in the tunnels and nests of other insects in wood and shoots with a soft core). Prey: various small Biology: Nests usually in dense sandy soil. Prey: spiders, mostly juvenile. Predators: emerald wasps dipterans Anthomyiidae, Calliphoridae, Muscidae, Trichrysis cyanea. Sacrophagidae, Syrphidae, Tabanidae, and Tachinidae. Predators: velvet ants Smicromyrme rufipes (Fabricius, Trypoxylon minus de Beaumont, 1945 1787) (Mutillidae) and parasitic flies Macronychia Literature: Mokrousov et al. 2013. griseola (Fallen, 1820), Metopia argyrocephala Material: RM: 1 male, 12.vi.2016, MSNR, quar. 413 (Meigen, 1824), Sphecapata conica Fallen, 1810, and (54045’12”N & 43016’54”E). Paramacronychia (Sarcophagidae). General distribution: Russia (the European part from the centre till Crimea and the east and Altai), Europe, ? Mellinus crabroneus (Thunberg, 1791) Turkey, and Kazakhstan (Antropov 2017b). Literature: Plavilshchikov 1964 (as Mellinus sabulosus F.) Biology: Nests in hollow stems or abandoned General distribution: Russia (the European part from tunnels of xylophagous insects in wood. Prey: juvenile the northwest to the southeast, Urals, Altai, eastern spiders Entelecara and Lepthyphantes [Linyphiidae], as Siberia, Amur region, and Primorsky Krai), Europe, well as Parasteatoda tepidariorum (C.L. Koch, 1841) and Turkey, Kazakhstan, Kyrgyzstan, Mongolia, northern Theridion varians Hahn, 1833 (Theridiidae). Predators: China, and Korea (Nemkov 2017d). Melittobia acasta chalcid wasps (Walker, 1839), Biology: Nests in sandy soil, often in groups along Tetrastichus (Eulophidae) and Eurytoma (Eurytomidae), the banks of water bodies. Prey: flies Anthomyiidae, ichneumons Ephialtes, Peritous (Ichneumonidae), Muscidae, and Syrphidae. and parasitic flies Cacoxenus indagator Loew, 1858 Remark: Confirmation of the species in Mordovia (Drosophilidae) (Olberg 1959). requires more material. Remark: The species is confirmed in MSNR.

Subfamily Dinetinae Subfamily Pemphredoninae Dinetus pictus (Fabricius, 1793) Tribe Pemphredonini Literature: Mokrousov et al. 2013. Diodontus luperus Shuckard, 1837 Material: RM: 1 male, 11.vii.2015, MSNR, quar. 142 Literature: Mokrousov et al. 2013. (54051’12”N & 43009’21”E). General distribution: Russia (the European part from General distribution: Russia (the European part the centre till Crimea and northern Caucasus, Urals, from the north to northern Caucasus and Altai), Europe, Altai, and eastern Siberia), Europe, northern Africa, Turkey, and Kazakhstan (Nemkov 2017c). Turkey, Kazakhstan, and Mongolia (Mokrousov 2017). Biology: Nests in the sandy soil. Prey: nymphs of Biology: Nests in dense soil on precipices and steep heteropterans Nabidae (mainly, Nabis) and Lygaeidae. slopes. Prey: nymphs and imagos of aphids. This species Pre-dators: emerald wasps Chrysis bicolor Lepeletier, kills victims by clenching its mandibles (Edwards & Telfer

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2002). from the north till Crimea and northern Caucasus), Europe, northern Africa, Turkey, Kazakhstan, and Japan * Diodontus medius Dahlbom, 1844 (Honshu) (Mokrousov 2017). Material: RM: 2 female, 14.vi.2016, MSNR, quar. 384 Biology: Nests in existing tubular cavities (in tunnels (54045’33”N & 43012’44”E); 1 female, 11.vi.2016, quar. of xylophagous insects in wood, in dry shoots of plants 389 (54045’50”N & 43017’50”E); 1 female, 1.vii.2016, with a soft core, and in hollow stalks of reeds). Prey: quar. 379 (54045’17”N & 43007’51”E); 1 female, nymphs and imagos of aphids. This species kills victims 9.vi.2016, quar. 358 (54046’05”N & 43011’44”E); 1 by clenching its mandibles, but it also often steals aphids female, 3.vii.2016, quar. 338 (54047’06”N & 43019’40”E); from the nests of other wasps. Predators: emerald 3 female, 13.vii.2017, quar. 319 (54048’09”N & wasps Omalus aeneus (Fabricius, 1787), Pseudomalus 43028’25”E). auratus (Linnaeus, 1758), and P. violaceus (Scopoli, General distribution: Russia (the European part 1763) (Chrysididae). from the north till the centre and the east, Urals, Tomsk region, Altai, Irkutsk, Amur and Magadan regions, ** Passaloecus gracilis (Curtis, 1834) Kamchatka, Primorsky Krai, and Sakhalin), Europe, and Material: RM: 1 female, 14.vi.2013, MSNR, Pushta Turkey (Mokrousov 2017). (54042’50”N & 43013’32”E). Biology: Nests in soil. Prey: nymphs and imagos General distribution: Russia (the European part from of aphids. This species kills victims by clenching its the centre till Crimea and northern Caucasus, Urals, mandibles. and Altai), Europe, northern Africa, Turkey, Abkhazia, Kazakhstan, Uzbekistan, Iran, and North America Diodontus minutus (Fabricius, 1793) (Mokrousov 2017). Literature: Mokrousov et al. 2013. Biology: Nests in existing tubular cavities (in tunnels General distribution: Russia (the European part from of xylophagous insects in wood, in dry shoots of plants the northwest till Crimea and northern Caucasus, Urals, with a soft core, and in galls of gallflies). Prey: nymphs Siberia, and Primorsky Krai), Europe, northern Africa, and imagos of Rhynchota (Sternorrhyncha) (Edwards & Israel, Kuwait, Turkey, Syria, Kazakhstan, central Asia, Telfer 2002; Giovanni et al. 2017). Predators: emerald Iraq, Iran, Mongolia, northern China, Korea, and North wasps Omalus aeneus (Fabricius, 1787), O. biaccinctus America (Mokrousov 2017). (du Buysson, 1892), and Pseudomalus auratus (Linnaeus, Biology: Nests in dense soil, often within the walls 1758) (Chrysididae). of forest cliffs. Prey: nymphs and imagos of aphids (Edwards & Telfer 2002). This species kills victims by Pemphredon fabricii (M. Müller, 1911) clenching its mandibles. Predators: emerald wasps Literature: Mokrousov et al. 2013. Chrysis lanceolata and C. leachii and velvet ants General distribution: Russia (the European part Myrmosa atra Panzer, 1801 and Smicromyrme sicanus from the centre to northern Caucasus), south of Europe, (De Stefani, 1887). Turkey, Kazakhstan, and Japan (Honshu) (Mokrousov 2017). Passaloecus borealis Dahlbom, 1844 Biology: Nests in rotten wood, in tunnels of Literature: Mokrousov et al. 2013. xylophagous insects, sometimes in hollow stems of General distribution: Russia (the European part plants or in shoots with a soft core. Prey: various aphids from the northwest to the centre, Altai, Irkutsk and (Bogusch et al. 2015). Magadan regions, Khabarovsk Krai and Primorsky Krai, and Sakhalin), Europe, Kazakhstan, and North America Pemphredon inornata Say, 1824 (Mokrousov 2017). Literature: Mokrousov et al. 2013. Biology: Nests in existing tubular cavities, usually Material: RM: 1 female, 15.viii.2015, Temnikov in tunnels of Anobiidae in wood. Prey: nymphs and Distr., Purdoshki (54040’04”N & 43032’33”E). imagos of aphids. This species kills victims by clenching General distribution: Russia (the European part its mandibles. from the northwest till Crimea and northern Caucasus, Urals, Siberia, Amur and Magadan regions, Kamchatka, Passaloecus corniger Shuckard, 1837 Khabarovsk Krai and Primorsky Krai, Sakhalin, and Kuril Literature: Ruchin & Antropov 2016. Islands), Europe, northern Africa, Turkey, Abkhazia, General distribution: Russia (the European part Kazakhstan, central Asia, Mongolia, northern China,

13238 Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov

Korea, Japan (Hokkaido and Honshu), North America from the northwest to northern Caucasus, Urals, (Mokrousov 2017). Siberia, Magadan region, Kamchatka, Khabarovsk Krai Biology: Nests in rotten wood, in tunnels of and Primorsky Krai, Sakhalin, and Kuril Islands), Europe, xylophagous insects, sometimes in hollow stems of Turkey, Kazakhstan, Kyrgyzstan, Uzbekistan, Mongolia, plants, in shoots with a soft core, or in galls of fliesLipara northern China, Korea, Japan (Hokkaido and Honshu), (Chloropidae). Prey: the most common and accessible and North America (Dollfuss 1995; Mokrousov 2017). aphids in the biotope. This species paralyzes prey with Biology: Nests in rotten wood, in tunnels of its sting or kills it by clenching its mandibles. Predators: xylophagous insects, sometimes in hollow stems of emerald wasps Pseudomalus auratus (Linnaeus, 1758) plants, in shoots with a soft core, or in galls of flies (Chrysididae). Lipara (Chloropidae). Prey: the most common and Remark: The species is confirmed in the territories accessible aphids in the biotope. This species paralyzes adjacent to MSNR. prey with its sting or kills it by clenching its mandibles. Predators: emerald wasps Pseudomalus triangulifer Pemphredon lethifer (Shuckard, 1837) (Abeille de Perrin, 1877) and P. violaceus (Scopoli, 1763) Literature: Ruchin & Antropov 2016. (Chrysididae). General distribution: Russia (the European part from the north till Crimea and northern Caucasus, Urals, ** Pemphredon rugifer (Dahlbom 1844) Novosibirsk region, Altai, Amur region, Khabarovsk Krai Material: RM: MSNR, Pushta (54042’50”N & and Primorsky Krai, Sakhalin, and Kuril Islands), Europe, 43013’32”E), 17.vi.2016, 1 female. northern Africa, Israel, Turkey, Abkhazia, Azerbaijan, General distribution: Russia (the European Kazakhstan, Kyrgyzstan, Uzbekistan, Iraq, Iran, part from the centre to northern Caucasus, Urals, Afghanistan, Mongolia, China, Korea, Japan (Hokkaido Novosibirsk region, Altai, Krasnoyarsk territory, and and Honshu), and North America (Mokrousov 2017). Sakhalin), Europe, Turkey, Kazakhstan, southwestern Biology: Nests in rotten wood, in tunnels of and northeastern China, Korea, Japan (Honshu), and xylophagous insects, sometimes in hollow stems of North America (Dollfuss 1995; Mokrousov 2017). plants, in shoots with a soft core, or in galls of flies Biology: Nests in rotten wood, in tunnels of Lipara (Chloropidae). Prey: the most common and xylophagous insects. Prey: the most common and accessible aphids in the biotope. This species paralyzes accessible aphids in the biotope. This species paralyzes prey with its sting or kills it by clenching its mandibles. prey with its sting or kills it by clenching its mandibles. Predators: emerald wasps Omalus aeneus (Fabricius, Predators: emerald wasps Trichrysis cyanea. 1787) (Chrysididae). Remark: Confirmation of the species in Mordovia requires more material. Spilomena troglodytes (Vander Linden, 1829) Literature: Mokrousov et al. 2013. Pemphredon lugens Dahlbom, 1843 General distribution: Russia (the European part from Literature: Mokrousov et al. 2009; Feoktistov 2011. the northwest till Crimea and northern Caucasus, Urals, General distribution: Russia (the European part and Primorsky Krai), Europe, northern Africa, Israel, from the northwest to the east, Urals, Altai, Krasnoyarsk Turkey, Abkhazia, Georgia, Kazakhstan, Uzbekistan, Krai, and Irkutsk and Magadan regions), Europe, Turkey, Kyrgyzstan, Tajikistan, and North America (Mokrousov and Kazakhstan (Dollfuss 1995; Mokrousov 2017). 2017). Biology: Nests in rotten wood, in tunnels of Biology: Nests in tunnels of xylophagous insects in xylophagous insects, sometimes in hollow stems of wood, hollow stems of reeds, or shoots with a soft core. plants, in shoots with a soft core, or in galls of fliesLipara Prey: nymphs of thrips. (Chloropidae). Prey: the most common and accessible aphids in the biotope. This species paralyzes prey with Stigmus pendulus Panzer, 1804 its sting or kills it by clenching its mandibles. Literature: Ruchin & Antropov 2016. General distribution: Russia (the European part from Pemphredon lugubris (Fabricius, 1793) the centre to northern Caucasus and Urals) and Europe Literature: Mokrousov et al. 2009; Feoktistov 2011; (Mokrousov 2017). Ruchin & Antropov 2014. Biology: Nests in tunnels of xylophagous insects in General distribution: Russia (the European part wood or in shoots with a soft core. Prey: aphids and,

Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 13239 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov probably, jumping plant lice. and in stems of plants, as well as in soil. Prey: small leafhoppers. Tribe Psenini Remark: The species is confirmed in MSNRand its Mimesa equestris (Fabricius, 1804) adjacent territories. Literature: Mokrousov et al. 2013. Material: RM: 1 female, 10.viii.2016, MSNR, quar. Mimumesa beaumonti (van Lith, 1949) 440 (54043’56”N & 43013’15”E). Literature: Mokrousov et al. 2013. General distribution: Russia (the European part from Material: RM: 1 female, 10.viii.2016, MSNR, quar. the northwest to the centre and the east, Urals, Siberia, 440 (54043’56”N & 43013’15”E). Magadan region, Khabarovsk Krai and Primorsky Krai, General distribution: Russia (the European part and Sakhalin), Europe, Turkey, Kazakhstan, and Korea from the northwest to the centre and the east, Urals, (Mokrousov 2017). Tomsk and Irkutsk regions, and Primorsky Krai), south of Biology: Nests in sandy soil, often in groups. Prey: Europe, and Kazakhstan (Mokrousov 2017). nymphs and imagos of Auchenorrhyncha. Predators: Biology: Nests in rotten wood and in abandoned emerald wasps Elampus panzeri (Fabricius, 1804) tunnels of xylophagous insects. Prey: small leafhoppers. (Crysididae). Remark: The species is confirmed in MSNR. Remark: The species is confirmed in MSNR. Mimumesa dahlbomi (Wesmael, 1852) Mimesa lutaria (Fabricius, 1787) Literature: Mokrousov et al. 2013. Literature: Redikortsev 1938 (as Psen shuckardi Material: RM: 2 female, 17.vii.2016, MSNR, quar. Dalb.); Plavilshchikov 1964 (as Psen shuckardi Wesm.). 330 (54046’44”N & 43011’09”E); 1 male, 29.v.2016, quar. Material: RM: 1 female, 7.vii.2016, MSNR, quar. 361 115 (54051’46”N & 43009’26”E); 1 male, 18.vii.2016, (54046’17”N & 43015’47”E). quar. 303 (54047’17”N & 43012’32”E). General distribution: Russia (the European part from General distribution: Russia (the European part from the northwest to the centre and the east, Urals, Altai, the northwest till Crimea and northern Caucasus, Urals, eastern Siberia, Amur and Magadan regions, Kamchatka, Altai, Irkutsk, Amur and Magadan regions, Kamchatka, Khabarovsk Krai and Primorsky Krai, Sakhalin, and Khabarovsk Krai and Primorsky Krai, Sakhalin, and Kuril Islands), Europe, Turkey, Kazakhstan, Uzbekistan, Kuril Islands), Europe, Turkey, Kazakhstan, Kyrgyzstan, Kyrgyzstan, Mongolia, northeastern China, Korea, Japan Mongolia, China, Korea, and Japan (Honshu) (Mokrousov (Hokkaido, Honshu, and Shikoku), and North America 2017). (Mokrousov 2017). Biology: Nests in rotten wood and in abandoned Biology: Nests in sandy soil, often in groups. Prey: tunnels of xylophagous insects. Prey” small leafhoppers nymphs and imagos of Auchenorrhyncha. Predators: Delphacidae, Fulgoridae, and Jassidae. emerald wasps Elampus panzeri (Fabricius, 1804) Remark: The species is confirmed in MSNR. (Crysididae). Remark: The species is confirmed in MSNR. Mimumesa unicolor (Vander Linden, 1829) Literature: Redikortsev 1938 (as Psen unicolor Mimumesa atratina (F. Morawitz, 1891) Wesm.); Plavilshchikov 1964 (as Psen unicolor Wesm.); Literature: Mokrousov et al. 2013; Ruchin & Mokrousov et al. 2013. Antropov 2014. Material: RM: 1 female, 23.vii.2016, Elniki Distr., Material: RM: 2 female, 13.vii.2017, MSNR, quar. Malye Mordovskie Poshaty (54041’00”N & 43043’50”E). 319 (54048’09”N & 43028’25”E); 1 female, 3.vi.2015, General distribution: Russia (the European part from Pushta (54042’50”N & 43013’32”E). the north to northern Caucasus, Urals, and Siberia), General distribution: Russia (the European part Europe, Israel, Turkey, Azerbaijan, Kazakhstan, central from the northwest to northern Caucasus, Urals, Asia, Iraq, Afghanistan, Pakistan, and China (Mokrousov Siberia, Magadan region, Kamchatka, Khabarovsk Krai 2017). and Primorsky Krai, Sakhalin, and Kuril Islands), Europe, Biology: Nests in soil, on clay cliffs or steep Turkey, Kazakhstan, Kyrgyzstan, Mongolia, northern bare slopes. Prey: small leafstones (Rhynchota and China, Korea, Japan (Honshu and Kyushu), and North Auchenorrhyncha) (Giovanni et al. 2017). Predators: America (Mokrousov 2017). emerald wasps Chrysis succincta Linnaeus, 1767, Biology: Nests in existing tubular cavities in wood Hedychridium ardens (Coquebert, 1801), Holopyga

13240 Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov gloriosa (Fabricius, 1793), and Pseudomalus auratus adjacent territories. (Linnaeus, 1758). Remark: The species is confirmed in the territories Psenulus pallipes (Panzer, 1798) adjacent to MSNR. Literature: Redikortsev 1938 ; Plavilshchikov 1964. Material: RM: 4 female, 17.vii.2016, MSNR, quar. 330 ? Psen ater (Olivier, 1792) (54046’44”N & 43011’09”E); 1 female, 21.vi.2015, quar. Literature: Redikortsev 1938 (as Psen ater Panz.); 381 (54045’17”N & 43009’52”E); 1 male, 12.vi.2016, quar. Plavilshchikov 1964 (as Psen ater F.). 430 (54044’34”N & 43016’00”E); 1 female, 13.vii.2017, General distribution: Russia (the European part quar. 347 (54047’34”N & 43028’31”E); 1 female, from the centre till Crimea, Urals, Altai, eastern Siberia, 19.vii.2016, cord. Podrubnyi (54047’51”N & 43008’48”E). Amur region, Khabarovsk Krai and Primorsky Krai, and General distribution: Russia (the European part Kuril Islands), Europe, Turkey, Kazakhstan, Uzbekistan, from the north till Crimea and northern Caucasus, Mongolia, northern China, Korea, and Japan (Hokkaido, Urals, Tomsk region, Altai, Irkutsk and Amur regions, Honshu, Shikoku, and Kyushu) (Mokrousov 2017). Kamchatka, Khabarovsk Krai and Primorsky Krai, Biology: Nests in sandy or clayey soil. Prey: Sakhalin, and Kuril Islands), Europe, Abkhazia, Armenia, leafstones. Predators: velvet ants Mutilla (Mutillidae) Turkey, Iran, Kazakhstan, northern and southwestern and ichneumons Perithous mediator (Fabricius, 1804) China, Korea, Japan (Hokkaido and Honshu), and North (Ichneumonidae). America (Mokrousov 2017). Remark: Confirmation of the species in Mordovia Biology: Nests in the existing tunnels of xylophagous requires more material. insects in wood and in dry hollow or soft-core plant stems. Prey: nymphs and imagos of aphids. Predators: emerald wasps of the genera Elampus, Omalus, Philoctetes, and Pseneo exaratus (Eversmann, 1849) Pseudomalus (Chrysididae) and ichneumons Perithous Literature: Ruchin & Antropov 2016. divinator (Rossi, 1790) and Perithous mediator (Fabricius, General distribution: Russia (the European part from 1804) (Ichneumonidae). the centre till Crimea, Urals, western Siberia, Irkutsk and Remark: The species is confirmed in MSNR. Amur regions, and Primorsky Krai), south of Europe, Abkhazia, Georgia, Kazakhstan, India, southeastern Subfamily Philanthinae China, Korea, and Japan (Hokkaido, Honshu, Shikoku, Tribe Cercerini and Ryukyu) (Mokrousov 2017). Cerceris arenaria (Linnaeus, 1758) Literature: Redikortsev 1938; Plavilshchikov 1964; Psenulus fuscipennis (Dahlbom, 1843) Mokrousov et al. 2013; Ruchin & Antropov 2014. Literature: Mokrousov et al. 2013. Material: RM: 1 male, 19.vii.2015, Elniki Distr., Malye Material: RM: 2 female, 5.vi.2015, MSNR, quar. 434 Mordovskie Poshaty (54041’00”N & 43043’50”E). 0 0 (54 45’04”N & 43 19’02”E); 1 male, 18.vii.2016, quar. General distribution: Russia (the European part 0 0 278 (54 47’51”N & 43 12’27”E); 1 female, 28.vi.2015, from the northwest till Crimea and northern Caucasus, 0 0 cord. Inorski (54 44’15”N & 43 08’53”E); 1 female, Urals, Siberia, Amur region, and Primorsky Krai), Europe, 0 1 male, 25.v.2014, cord. Podrubnyi (54 47’51”N & northern Africa, Israel, Turkey, Caucasus, Kazakhstan, 0 43 08’48”E); 2 female, 16.vii.2016, 17.vii.2017, Pushta central Asia, Iran, Mongolia, northeastern China, Korea, 0 0 (54 42’50”N & 43 13’32”E). and Japan (Honshu and Ryukyu) (Danilov & Mokrousov General distribution: Russia (the European part from 2017). the north till Crimea and northern Caucasus, Urals, and Biology: Nests in dense clay or sandy soil. Prey: Primorsky Krai), Europe, Georgia, Turkey, India, Korea, snout beetles, more often Brachyderes, Curculio, and Japan (Hokkaido and Honshu) (Mokrousov 2017). Hylobius, Otiorhynchus, Pissodes, Sitona, Strophosomus, Biology: Nests in tubular cavities in wood, in hollow and Tanymecus and sometimesBromius (Curculionidae) or soft-core stems of plants, and in old galls of flies (Polidori et al. 2005). Predators: emerald wasps Chrysis Lipara (Chloropidae). Prey: nymphs and imagos of ignita (Linnaeus, 1758), C. sexdentata Christ, 1791, aphids. Predators: Omalus sculpticollis, Melittobia Hedychrum niemelai Linsenmaier, 1959, H. nobile acasta, Perithous septemcinctorius, and Trichodes (Scopoli, 1763), H. rutilans Dahlbom, 1854, and H. simile apiarius (Martynova & Fateryga 2014). Mocsáry, 1889 (Chrysididae), velvet ants Smicromyrme Remark: The species is confirmed in MSNRand sicanus (De Stefani, 1887) (Mutillidae), parasitic flies

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Metopia leucocephala (Rossi, 1790), Miltogramma Elniki Distr., Novye Shaly (54042’13”N & 43038’25”E). punctatum Meigen, 1824 (Sarcophagidae), and General distribution: Russia (the European part Leucophora cinerea Robineau-Desvoidy, 1830 from the northwest till Crimea and northern Caucasus, (Anthomyiidae), as well as ants Tetramorium caespitum Urals, and Siberia), Europe, northern Africa, Turkey, (Linnaeus, 1758) (Formicidae), ticks of the genera Vidia Azerbaijan, and Kazakhstan (Danilov & Mokrousov (Winterschmidtiidae), and nematodas (Polidori et al. 2017). 2010). Biology: Nests in dense soil. Prey: small snout beetles Apion, Meligethes, Sitona, Strophosomus, and Tychius Cerceris interrupta (Panzer, 1799) (Curculionidae). Predators: emerald wasps Hedychrum Literature: Ruchin et al. 2009. niemelai Linsenmaier, 1959 (Chrysididae). Material: RM: 1 female, 23.vii.2016, Elniki Distr., Malye Mordovskie Poshaty (54041’00”N & 43043’50”E); Cerceris ruficornis (Fabricius, 1793) NNR: 1 ex., 16.viii.2014, Sarov (54055’08” N 43017’40”E), Literature: Ruchin & Antropov 2014. coll. P.V. Cherenkov (Image 3). Material: RM: 1 male, 11.vii.2015, MSNR, quar. 115 General distribution: Russia (the European part (54051’46”N & 43009’26”E); 1 female, 1 male, 11.vii.2015, from the centre till Crimea and northern Caucasus, quar. 142 (54051’12”N & 43009’21”E); 1 female, Urals, Novosibirsk region, Altai, and Krasnoyarsk Krai), 11.vii.2015, quar. 301 (54047’19”N & 43010’36”E); 1 south of Europe, Turkey, and Kazakhstan (Danilov & male, 18.vii.2016, quar. 331 (54046’43”N & 43012’37”E); Mokrousov 2017). 1 male, 1.vii.2016, quar. 379 (54045’17”N & 43007’51”E); Biology: Nests in dense soil. Prey: snout beetles 1 male, 4.vii.2017, quar. 276 (54047’45”N & 43010’28”E). (Curculionidae) and rarely seed beetles (Bruchidae). General distribution: Russia (the European part Predators: emerald wasps Hedychrum chalybaeum from the north till Crimea and the east, Urals, Siberia, Dahlbom, 1854 (Chrysididae). Khabarovsk Krai, and Sakhalin), Europe, northern Africa, Turkey, Kazakhstan, Uzbekistan, Kyrgyzstan, Taldikistan, Cerceris quadrifasciata (Panzer, 1799) Iran, northern China, and Korea (Danilov & Mokrousov Literature: Mokrousov et al. 2013. 2017). Material: RM: 1 female, 19.vii.2016, MSNR, quar. Biology: Nests in dense soil. Prey: snout beetles 330 (54046’44”N & 43011’09”E); NNR: 1 ex., 19.vii.2014, Apion, Otiorrhynchus, Sitona, and Strophosomus Voznesenskoe Distr., Alamasovo (54054’03”N & (Curculionidae) as well as leaf beetles Longitarsus and 43008’56”E), coll. P.V. Cherenkov. Bromius (Chrysomelidae), and, probably, heteropterans. General distribution: Russia (the European part from Predators: emerald wasps Hedychrum gerstaeckeri the northwest till Crimea and northern Caucasus, Urals, Chevrier, 1869, H. niemelai Linsenmaier, 1959, and H. Siberia, Amur region, Khabarovsk Krai and Primorsky nobile (Scopoli, 1763) (Chrysididae). Krai, and Sakhalin), Europe, Turkey, Armenia, Georgia, Remark: The species is confirmed in MSNR. Abkhazia, Kazakhstan, northeastern China, and Korea (Danilov & Mokrousov 2017). Cerceris rybyensis (Linnaeus, 1771) Biology: Nests in dense soil, often in large groups. Literature: Mokrousov et al. 2009, 2013; Feoktistov Prey: snout beetles Anthonomus, Dorytomus, 2011; Ruchin & Antropov 2014. Otiorrhynchus, Phyllobius, Polydrosus,and Strophosomus Material: RM: 1 male, 3.vii.2016, MSNR, quar. (Curculionidae). Predators: emerald wasps Hedychrum 429 (54044’31”N & 43014’59”E); 1 male, 24.vi.2014, niemelai Linsenmaier, 1959 H. nobile (Scopoli, 1763) and Pushta (54042’50”N & 43013’32”E); 2 male, 19.vii.2015, Holopyga chrysonota (Förster, 1853) (Chrysididae). Elniki Distr., Malye Mordovskie Poshaty (54041’00”N Remark: The species is confirmed in MSNR. & 43043’50”E); 1 female, 22.vii.2017, Novoyamskaya Sloboda (54044’24”N & 43042’59”E); 1 male, 22.vii.2017, Cerceris quinquefasciata (Rossi, 1792) Novye Shaly (54042’13”N & 43038’25”E). Literature: Redikortsev 1938; Plavilshchikov 1964; General distribution: Russia (the European part Ruchin & Antropov 2014, 2016. from the northwest till Crimea and northern Caucasus, Material: RM: 1 male, 19.vii.2016, MSNR, quar. 330 Urals, Siberia, Primorye, and Sakhalin), Europe, northern (54046’44”N & 43011’09”E); 1 male, 18.vii.2016, quar. Africa, Turkey, Georgia, Azerbaijan, Syria, Kazakhstan, 331 (54046’43”N & 43012’37”E); 1 female, 22.vii.2017, Uzbekistan, Turkmenistan, Tajikistan, Iran, India, Korea,

13242 Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov

A B

C D

E F

G H

Image 2. Images of wasps (© P.V. Cherenkov). A - Hedychrum longicolle (Sarov, 18.vi.2015), B - Trichrysis cyanea (Voznesenskoe Distr., Alamasovo, 14.vi.2015), C - Scolia hirta (Sarov, 8.viii.2014), D - Vespa crabro with prey Vespula vulgaris (Sarov, 6.viii.2014), E - Vespula germanica (Sarov, 31.viii.2014), F - Bembix rostrata (Sarov, 8.viii.2014), G - Ectemnius borealis (Sarov, 29.v.2015), H - Cerceris interrupta (Sarov, 16.viii.2014).

Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 13243 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov

A B

D C

E F

Image 3. Images of wasps. A - Ancistrocerus trifasciatus (Voznesenskoe Distr., Alamasovo, 22.vii.2017, © P.V. Cherenkov), B - Polistes nimpha (cord. Inorski, 6.ix.2014, © G.B. Semishin), C - Podalonia hirsuta (Voznesenskoe Distr., Alamasovo, 16.vii.2017, © P.V. Cherenkov), D - Sphex funerarius (Sarov, 5.viii.2014, © P.V. Cherenkov), E - Gorytes laticinctus (cord. Inorski, 2.vi.2017, © G.B. Semishin), F - Bembix rostrata (cord. Inorski, 27.vii.2017, © G.B. Semishin).

and Japan (Honshu) (Danilov & Mokrousov 2017). niemelai Linsenmaier, 1959, H. nobile (Scopoli, 1763), Biology: Nests in dense sandy or clayey soil, often and Chrysis ignita (Linnaeus, 1758) (Chrysididae), in groups. Prey: bees of genera Halictus, Andrena, velvet ants Paramyrmosa brunnipes (Lepeletier, 1845) Panurgus, Prosopis, and Sphecodes. Predators: emerald (Mutillidae), and parasitic flies Metopia leucocephala wasps Hedychrum gerstaeckeri Chevrier, 1869, H. (Rossi, 1790) and Pterella grisea (Meigen, 1824)

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(Sarcophagidae). Paeudogonalis hahni Spin.). General distribution: Russia (the European part, Philanthus triangulum (Fabricius, 1775) Altai, Irkutsk region, Transbaikalia, Amur region, and Literature: Redikortsev 1938; Plavilshchikov 1964; Primorsky Krai), Europe, Ukraine, Turkey, Kazakhstan, Mokrousov et al. 2009, 2013; Ruchin & Antropov 2014. Iran, Mongolia, and northern China (Schnee 2011). Material: RM: 1 male, 11.vii.2015, MSNR, quar. 301 Biology: Parasites of larvae and caterpillars of (54047’19”N & 43010’36”E); 1 female, 24.viii.2017, quar. butterflies (Schnee 2011). 80 (54053’11”N & 43030’22”E); 1 female, 24.viii.2017, quar. 85 (54053’29”N & 43035’00”E); 1 female, 24.viii.2017, quar. 86 (54053’38”N & 43035’59”E); RESULTS AND DISCUSSION 20.vi.2015, 1 male, Pushta (54042’50”N & 43013’32”E); 24.vii.2016, 1 male, Temnikov Distr., Polyanki Thus, we summarized data on 249 vespine species (54044’27”N & 43029’04”E); NNR: 23.vii.2016, 1 ex., of 11 families from two regions of Mordovia (including Sarov (54055’08” N 43017’40”E), coll. P.V. Cherenkov. MSNR) and three districts of Nizhny Novgorod region General distribution: Russia (the European part (including the city of Sarov). Of these, 31 species are new from the northwest till Crimea and northern Caucasus, to the Republic of Mordovia. In recent years, the fauna Urals, and western Siberia), Europe, Africa, Israel, of Nizhni Novgorod region was studied quite intensively (Mokrousov 2011; Nemkov 2016). The areas adjacent to Jordan, Yemen, Saudi Arabia, Turkey, Abkhazia, Georgia, the reserve areas of Nizhny Novgorod region, however, Azerbaijan, Kazakhstan, central Asia, Iraq, Iran, and were so far poorly explored. We provide information on Afghanistan (Danilov & Mokrousov 2017). 30 species of wasps occurring in these areas, of which Biology: Nests in dense sandy and sandy loam soil, two species were not yet recorded in the reserve. usually in groups. Prey: honey bees Apis mellifera As of 2015, 179 species of wasps were recorded in Linnaeus, 1758 or, as an exception, solitary bees, for MSNR (Redikortsev 1938; Plavilshchikov 1964; Ruchin et example, Andrena and Halictus (Rathmayer, 1962). al. 2009; Ruchin 2011, 2015; Mokrousov et al. 2009, 2013; Predators: emerald wasps Chrysis ignita (Linnaeus, Ruchin, Antropov, 2014, 2016). At present, the fauna of 1758), Hedychrum gerstaeckeri Chtvrier, 1869, H. separate families of wasps in MSNR is represented by rutilans Dahlbom, 1854, Pseudomalus auratus (Linnaeus, 220 species: one species in Bethylidae, 31 in Chrysididae, 1758) (Chrysididae), velvet ants Dasylabris maura two in Tiphiidae, two in Mutillidae, one in Scoliidae, 28 (Linnaeus, 1758) (Mutillidae), parasitic flies Senotainia in Pompilidae, 37 in Vespidae, nine in Sphecidae, 108 in albifrons Phylloteles pictipennis (Rondani, 1859), Loew, Crabronidae, and one in Trigonalyidae. 1824, Metopia leucocephala (Rossi, 1790), Sphecapata Unfortunately, in the 20th Century, the authors of conica Fallen, 1810 (Sarcophagidae), Physocephala publications on fauna of MSNR limited themselves chrysorrhoea (Meigen, 1824) and Ph. vittata (Fabricius, to bringing up short lists of wasps without specifying 1794) (Conopidae), and skin beetles Dermestes murinus important characteristics such as the place and date of Linnaeus, 1758 (Dermestidae) (Polidori et al. 2010). collection, as well as the sex and the number of material studied. This complicates the comparison of the species Philanthus venustus (Rossi, 1790) composition, biotopic confinement, and verification of Literature: Ruchin & Antropov 2014. finds in certain habitats. Therefore, a number of species General distribution: Russia (the European part are indicated so far only on the basis of these literary from the centre and east till Crimea and northern sources. Among them, in particular, are such species Caucasus and Urals), south of Europe, northern Africa, whose modern ranges include the territory of Mordovia: Israel, Turkey, Syria, Kazakhstan, and Mongolia (Danilov A. wesmaeli (Redikortsev, 1938), P. ater (Redikortsev, & Mokrousov 2017). 1938; Plavilshchikov, 1964), Ch. pumila, Ch. comparata, Remark: Confirmation of the species in the territories P. tydei, N. spinolae, M. crabroneus (Plavilshchikov, adjacent to MSNRrequires more material. 1964), A. terminata, C. ovalis, and T. fronticorne (Feoktistov 2011). Superfamily Trigonalyoidea At the same time, some of the species included in Family Trigonalyidae these lists, according to modern data, clearly do not Pseudogonalos hahnii (Spinola,1840) occur in the territory of Mordovia, since their range Literature: Ruchin et al. 2009; Feoktistov 2011 (as in the European part of Russia is limited to the south,

Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 13245 Wasp fauna of Mordovia State Nature Reserve and its surroundings Ruchin & Antropov including Crimea and northern Caucasus. Such species the MSNRlist of wasp fauna. include C. octomaculatus (Ghahari et al. 2014; Loktionov Of particular interest is S. deforme, an eastern Asian & Lelej 2017). It is most likely that these records were species that in recent years showed an expansion of based on incorrect definition of the source material. the range to the northwest (Rasplus 2010; Ćetković et Therefore, we excluded this species from the MSNR al. 2011). Judging by the many-year collections, there fauna list. is a noticeable increase in its population in Mordovia in In recent years, the list of wasps species recorded in comparison with the native species, S. destillatorium. MSNRand adjacent territories significantly expanded. Similarly, P. fastigiata, noted in Mordovia for the first Many species from the list, however, are still known time, which was previously known to the north no only by single specimens. To such species we refer, in further than the Rostov region (Savranskaya & Lidzhiev particular, C. megacephalus (Mokrousov et al. 2009), E. 2017), possibly also expands its area to the northwest. minor, G. lunatus, P. gussakowskiji, P. minuta, P. parvul, Additional material, however, is required to confirm P. schioedtei, A. concinnus, A. anceps, H. sanguinolentus, this. A. auctus, M. parvulus, D. lineata, N. dimidiatus, N. The rare species listed in the Red Book of the Russian fulvipes, C. annulipes, C. elongatulus, C. exiguus, O. Federation (2001) include P. grandior. Ten-year studies haemorrhoidalis, P. fabricii, S. troglodytes (Mokrousov in MSNRshow that the number of Parnopes is very et al. 2013), O. simillimus (Ruchin & Antropov 2014), low (Ruchin & Egorov 2017). Perhaps the overgrowth A. ichneumonideus, E. quadrifasciatus, G. laevipes, S. of glades and ravines with shrub vegetation and, as a gracilis, P. lethifer, S. pendulus, and P. exaratus (Ruchin consequence, degradation of host-wasps colonies, have a negative impact on this population (Orlovskytė et al. & Antropov 2016). Records of these species are 2010). A similar situation is observed for the Apollo undoubtful. Butterfly Parnassius apollo (Linnaeus, 1758), whose The most common in the reserve are species of habitat in the reserve is lost precisely because of the the families Chrysididae (Hedychrum nobile, Chrysis same reasons (Ruchin & Grishutkin 2018). In the case of bicolor, C. ignita, and Trichrysis cyanea), Tiphiidae Parnopes, however, the situation is not that bad, as very (Tiphia femorata), Pompilidae (Auplopus carbonarius, suitable habitats for host-wasps appeared during the Priocnemis exaltata, P. perturbator, and Anoplius fires of 2010. Another rare species that requires further viaticus), Vespidae (Ancistrocerus trifasciatus, Eumenes study is S. hirta. This species as well as P. fastigiata coronatus, E. pedunculatus, Symmorphus bifasciatus, S. gradually expand their range to the north (Ruchin & murarius, Polistes albellus, P. nimpha (dominant), Vespa Artaev 2016). crabro, Dolichovespula saxonica, Vespula germanica, A significant majority of the identified representatives V. vulgaris Ammophila pubescens, and ), and Sphecidae ( of the emerald wasp family are brood parasites of A. sabulosa, Podalonia hirsuta, Argogorytes mystaceus, digger wasps (Crabronidae) while some are brood Gorytes quadrifasciatus, G. quinquecinctus, Bembecinus parasites of the solitary Vespidae. We point out that tridens, Nysson spinosus, Crabro cribrarius, Ectemnius in the overwhelming majority of cases, the hosts of the borealis, E. continuus, E. lapidarius, Lestica clypeata emerald wasps were also revealed during the course of (dominant), and Cerceris quinquefasciata). research. The strength of brood parasite-host relations In a recent publication on European paper wasps indicates the stability of biocenoses in MSNR. (Polistes) of the gallicus-group (Neumeyer et al. Digger wasps (Sphecidae and Crabronidae) make 2015), the status of an independent species was given up an important part of the Aculeata fauna (53.2%), to Polistes albellus Giordani Soika, 1976, originally counting 117 species. In forest biotopes, the prevailing described as a subspecies of P. foederatus Kohl, 1898 species were compared to the terrestrial species and included in the list of lower synonyms P. gallicus (67.5%) as xylophile and rubicole (21.4%). Species of (Linnaeus, 1767). For a long time,P. albellus, distributed the genera Passaloecus and Trypoxylon make their nests from the centre and east of the European part and south in the wood in the finished galleries of wood-destroying of western Siberia of Russia to Mongolia, was confused insects while species of the genera Crossocerus and with similar species P. bischoffi Weyrauch, 1937 and P. Ectemnius complete the galleries in the wood with biglumis (Linnaeus, 1758), which in Russia are found their mouth organs. Almost all species nesting in the mainly in northern Caucasus. Therefore, we excluded soil choose open sites with sparse vegetation, more or the previously mentioned P. bischoffi and P. biglumis less flat (dry glades, roadsides and clearings, open areas (Mokrousov et al. 2013; Ruchin & Antropov 2014) from with sandy face, and slopes of small pits). Sand is a drier

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Table 1. Contribution of MSNR to the vespine diversity of Mordovia.

Vespine diversity of Number of species in Number of species in Families the reserve and the MSNR Mordovia (this study) region (in %) Bethylidae 1 1 100

Chrysididae 31 41 75.6

Dryinidae 0 1 0

Tiphiidae 2 5 40.0

Mutillidae 2 3 66.7

Scoliidae 1 2 50.0

Pompilidae 28 38 73.7

Vespidae 37 47 78.7

Sphecidae 9 12 75.0

Crabronidae 108 132 81.8

Trigonalyidae 1 1 100

Total 220 283 77.7

and generally more favorable nesting substrate than al. 2011; Ruchin et al. 2013; Chikhlyaev & Ruchin 2014; clayey soil. Therefore, the greatest species diversity of Legalov et al. 2014; Egorov & Shapovalov 2017; Chursina digger wasps is typical for the central and eastern parts & Ruchin 2018a, 2018b; Ruchin & Egorov 2018a, of the reserve, but not for the floodplain meadows of 2018b, 2018c, 2018d; Ruchin et al. 2018. 2019; Ruchin the western part where the grass cover is very dense. & Mikhailenko 2018; Tomaszewska et al. 2018). Such Some species of the Sphecidae family that were found an advantageous geographic location of the reserve in the adjacent territories were not recorded as part of at the edge of the forest-steppe and forest zone also the reserve fauna. This reduction is usually due to the significantly affects the species diversity and generally overgrowth of biotopes necessary for Sphecidae (Kula & contributes to the biodiversity of the vespine species in Tyrner 2003). the Republic of Mordivia Mordovia Republic (Table 1). Most of the identified species of Sphecidae prey Of the 283 species of wasps in Mordovia, 220 species on Lepidoptera, Hymenoptera, and Aranei. In the (77.7%) are found within MSNR. Crabronidae family, a significant number of species consume medium and small spiders, whose fauna in the reserve is rich enough (Mikhailov & Trushina 2013; REFERENCES Ruchin et al. 2013). Some Crabronidae hunt for Diptera, Aguiar, A.P., A.R. Deans, M.S. Engel, M. Forshage, J.T. Huber, J.T. Heteroptera, Homoptera (Cicadellidae, Psyllidae, Jennings, N.F. Johnson, A.S. Lelej, J.T. Longino, V. Lohrmann, I. Jassidae, and Fulgoridae), and Orthoptera. Mikó, M. Ohl, C. Rasmussen, A. Taeger & D.S.K.Yu (2013). Order MSNR is one of the oldest in Russia (Vargot 2016). Hymenoptera. Zootaxa 3703(1): 51–62; https://doi.org/10.11646/ zootaxa.3703.1.12 According to paleoecological reconstructions (Novenko Antropov, A.V. (2017a). 68. Family Vespidae. Subfamily Polistinae et al. 2017), since 5,000 years ago broad-leaved forests – Paper wasps, Subfamily Vespinae – Hornets and Jellow Jackets, prevailed in suitable ecotopes on the territory of the pp. 193–196. In: Belokobylskij, S.A. & A.S. Lelej (eds). Annotated Catalogue of the Hymenoptera of Russia, Vol. I. Symphyta and reserve. Significant territories were also covered with : Aculeata. Proceedings of the Zoological Institute of the pine forests, possibly with very little participation of Russian Academy of Sciences Supplement 6: 1–475. spruce. Pine forests grew on poor sandy soils and in Antropov, A.V. (2017b). 72. Family Crabronidae. Subfamily Crabroninae, pp. 225–245. In: Belokobylskij, S.A. & A.S. Lelej poorly drained habitats. This fully corresponds to the (eds). Annotated Catalogue of the Hymenoptera of Russia, Vol. I. location of mixed coniferous-broad-leaved forests Symphyta and Apocrita: Aculeata. Proceedings of the Zoological Institute of the Russian Academy of Sciences Supplement 6: 1–475. zone in the south. In addition, the territory refers Antropov, A.V. & M.V. Mokrousov (2016). New data on distribution of to landscapes of the marshy woodland type, which four species of the genus Trypoxylon (Hymenoptera: Crabronidae: are extrazonal boreal geosystems. Recent records of Trypoxylini) in European Russia. Russian Entomological Journal 25(3): 265–269. invertebrates from the territory of the reserve are very Blösch, M. (2000). Die Grabwespen Deutschlands. Lebensweise, interesting and confirm its uniqueness (Mokrousov et Verhalten, Verbreitung. Die Tierwelt Deutschlands 71, Goecke &

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Fauna Entomologica Annotated Catalogue of the Hymenoptera of Russia, Vol. I. Symphyta Scandinavica 30: 1–98. and Apocrita: Aculeata. Proceedings of the Zoological Institute of Orlovskytė, S., A. Budrienė & E. Budrys (2010). Check-list of cuckoo- the Russian Academy of Sciences Supplement 6: 246–252. wasps (Hymenoptera: Chrysididae) of Lithuania. New and Rare for Mokrousov, M.V. & A.S. Lelej (2017). 62. Family Scoliidae. 63. Lithuania Insect Species 22: 141–156. Family Tithiidae, pp149–150. In: Belokobylskij, S.A. & A.S. Lelej Peters, R.S., L. Krogmann, Ch. Mayer, A. Donath, S. Gunkel, K. (eds). Annotated Catalogue of the Hymenoptera of Russia, Vol. I. Meusemann, A. Kozlov, L. Podsiadlowski, M. Petersen, R. Lanfear, Symphyta and Apocrita: Aculeata. Proceedings of the Zoological P.A. Diez, J. Heraty, K.M. Kjer, S. Klopfstein, R. Meier, C. Polidori, Th. Institute of the Russian Academy of Sciences Supplement 6: 147– Schmitt, Sh. Liu, X. Zhou, T. Wappler, J. Rust, B. Misof & O. Niehuis 148. (2017). Evolutionary History of the Hymenoptera. Current Biology Mokrousov, M.V., A.V. Antropov, A.B. Ruchin & K.E. Bugayev (2009). 27(7): 1013–1018; https://doi.org/10.1016/j.cub.2017.01.027 Some information on fauna of wasps (Hymenoptera: Chrysididae, Plavilshchikov, N.N. (1964). A list of insect species found on the Tiphiidae, Pompilidae, Vespidae, Sphecidae, Crabronidae) of territory of the Mordovia State Nature Reserve. Proceedings of the Mordovia Reserve (based on collections). Proceedings of the Mordovia State Nature Reserve 2: 105–134 [in Russian]. Stavropol Branch of the Russian Entomological Society 5: 77–79 [in Polidori, C., S. Bevacqua & F. Andrietti (2010). Do digger wasps time Russian]. their provisioning activity to avoid cuckoo wasps (Hymenoptera: Mokrousov, M.V., A.B. Ruchin & L.V. Egorov (2013). The wasp Crabronidae and Chrysididae)? Acta Ethologica 13(1): 11–21; fauna of Mordovia State Nature Reserve and adjacent territories. https://doi.org/10.1007/s10211-010-0067-4 Proceedings of the Mordovia State Nature Reserve 11: 193–205 [in Polidori, C., R. Boesi, F. Isola & F. Andrietti (2005). Provisioning Russian]. patterns and choice of prey in the digger wasp Cerceris arenaria Murai, M.N. & Z.S. Amr (2011). Sphecidae and Crabronidae of the (Hymenoptera: Crabronidae): the role of prey size. European Al Thawrah Nature Reserve, northern Syria, with a checklist of the Journal of Entomology 102(4): 801–804; https://doi.org/10.14411/ species known from Syria (Insecta: Hymenoptera). Zoology in the eje.2005.111 Middle East 53: 107–120. Ponomarenko, N.G. & M. Olmi (2007). New records of Dryinidae Nemkov, P.G. (2008). Digger wasps of the genus Nysson Latreille (Hymenoptera: Chrysidoidea) from Russia, Ukraine, Moldova, (Hymenoptera, Crabronidae) of the Russian fauna. Euroasian Georgia, Armenia and Azerbaijan. Entomologist’s Gazette 58: 51– Entomological Journal 7(3): 217–221. 58. Nemkov, P.G. (2012). Digger wasps of the genus Bembecinus A. Costa, Rasplus, J.-Y. (2010). 14.61 – Sceliphron curvatum (Smith, 1870), 1859 (Hymenoptera, Crabronidae, Bembicinae) of the fauna of S. caementarium (Drury, 1773) and S. deforme (Smith, 1856) Russia and neighbouring countries. Far Eastern Entomologist 251: (Hymenoptera, Sphecidae). In: Roques, A. et al. (eds.). 1–11. Invasions in Europe. BioRisk 4(2): 982–983; https://doi.org/10.3897/ Nemkov, P.G. (2016). Digger wasps of the genus Bembix Fabricius, biorisk.4.69 1775 (Hymenoptera: Crabronidae, Bembicinae) of Russia and Rathmayer, W. (1962). Paralysis caused by the digger wasp Philanthus. adjacent territories. Far Eastern Entomologist 313: 1–34. Nature 196: 1149–1151. Nemkov, P.G. (2017a). Subfamily Astatinae, pp218–219. In: Redikortsev, V.V. (1938). Materials to the entomofauna of the Belokobylskij, S.A. & A.S. Lelej (eds). Annotated Catalogue of the Mordovia State Nature Reserve, pp137–146. In: Fauna of the Hymenoptera of Russia, Vol I. Symphyta and Apocrita: Aculeata. Mordovia State Nature Reserve. Committee of Nature Reserves Proceedings of the Zoological Institute of the Russian Academy of at the Presidium of the All-Union Central Executive Committee, Sciences Supplement 6: 1–475. Moscow, 153p. [in Russian]. Nemkov, P.G. (2017b). Subfamily Bembicinae, pp219–225. In: Red Data Book of Russian Federation (2001). . Astrel, Belokobylskij, S.A. & A.S. Lelej (eds). Annotated Catalogue of the Moscow, 862pp [in Russian]. Hymenoptera of Russia, Vol I. Symphyta and Apocrita: Aculeata. Rosa, P., A.S. Lelej, S.A. Belokobylskij, V.M. Loktionov & L.A. Zaytseva Proceedings of the Zoological Institute of the Russian Academy of (2017a). 60. Family Chrysididae – cuckoo wasps. In: Belokobylskij, Sciences Supplement 6: 1–475. S.A. & A.S. Lelej (eds). Annotated Catalogue of Hymenoptera of Nemkov, P.G. (2017c). 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Zoological Institute of the Russian Academy of Sciences Supplement Ruchin, A.B. & G.F. Grishutkin (2018). Biology and distribution of 6: 126–144. Parnassius apollo (Linnaeus, 1758) a rare species in Mordovia Rosa, P., A.S. Lelej, M.Yu. Proshchalykin, V.M. Loktionov & M.V. Republic, Russia. Biodiversitas 19(2): 431–436; https://doi. Mokrousov (2017b). New records of Chrysididae (Hymenoptera) org/10.13057/biodiv/d190210 from Russia with description of five new species. Far Eastern Ruchin, A.B. & D.K. Kurmaeva (2010). On rare insects of Entomologist 345: 1–33; https://doi.org/10.25221/fee.345.1 Mordovia included in the Red Book of the Russian Federation. Ruchin, A.B. (2011). First additional materials towards the Entomological Review 90(6): 712–717; https://doi.org/10.1134/ entomofauna of the Mordovia State Nature Reserve. Proceedings S0013873810060060 of the Mordovia State Nature Reserve 9: 150–182 [in Russian]. Ruchin, A.B. & N.А. Lengesova (2012). Preliminary information on the Ruchin, A.B. (2015). Second additional materials on the insect fauna fauna of Hymenoptera (Hymenoptera, Symphyta) of some specially of the Mordovia State Nature Reserve. Proceedings of the Mordovia protected natural territories of Mordovia Republic. Proceedings of State Nature Reserve 13: 351–398 [in Russian]. Khvalynsky National Park 4: 31–38 [in Russian]. Ruchin, A.B. & A.V. Antropov (2014). On the wasp fauna (Hymenoptera, Ruchin, A.B. & N.V. Makarkin (2017). Neuroptera and Raphidioptera Vespomorpha) of the Republic of Mordovia. Actual Problems of the in the Mordovia State Nature Reserve. Nature Conservation Humanities and Natural Sciences 3–1: 29–36 [in Russian]. Research 2(2): 38–46; https://doi.org/10.24189/ncr.2017.001 [in Ruchin, A.B. & A.V. Antropov (2016). New and interesting findings Russian] of wasps (Insecta, Hymenoptera) in the Mordovia Nature Reserve. Ruchin, A.B. & A.P. Mikhailenko (2018). Fauna of mantids and Proceedings of the Mordovia State Nature Reserve 16: 399–403 [in orthopterans (Insecta: Mantodea, Orthoptera) of the Mordovia State Nature Reserve, Russia. Biodiversitas 19(4): 1194–1206; Russian]. https://doi.org/10.13057/biodiv/d190403 Ruchin, A.B., Antropov, A.V. & A.A. Khapugin (2019). Distribution, Ruchin, A.B., M.K. Ryzhov, O.N. Artaev & A.A. Khapugin (2013). New abundance, and habitats of rare species Parnopes grandior (Pallas records of Argiope bruennichi (Scopoli, 1772) (Aranei: Araneidae) 1771) (Hymenoptera, Chrysididae) in Mordovia and adjacent from Mordovia and adjacent regions of Russia. Arthropoda Selecta regions, Russia. Biodiversitas 20(2): 303–310; https://doi. 22(4): 361–362. org/10.13057/biodiv/d200201 Ruchin, A.B. & S.V. Shibaev (2008). On the fauna of bees (Hymenoptera, Ruchin, A.B. & O.N. Artaev (2016). On expansion of the distribution Apoidea) of Mordovia, pp335–341. In: Modern Ecology - The Science range of some scoliid wasps (Scoliidae, Hymenoptera, Insecta) in of the 21st Century. Ryazan State University, Ryazan, 458pp [in the Middle Volga region. Research Journal of Pharmaceutical, Russian]. Biological and Chemical Sciences 7(3): 2110–2115. Ruchin, A.B. & V.A. Zryanin (2013). To the fauna of ants (Hymenoptera: Ruchin, A.B. & L.V. Egorov (2017). Overview of insect species included Formicidae) of the Republic of Mordovia, pp108–110. In: Ants and in the Red Data Book of Russian Federation in the Mordovia State Forest Protection. KMK Scientific Press Ltd., Moscow, 340pp [in Nature Reserve. Nature Conservation Research 2(1): 2–9; https:// Russian]. doi.org/10.24189/ncr.2017.016 [in Russian] Savranskaya, Zh.V. & O.N. Lidzhiev (2017). Materials on the fauna of Ruchin, A.B. & L.V. Egorov (2018a). Discovery of Allonyx road wasps (Hymenoptera, Pompilidae) of Kalmykia, pp34–36. In: quadrimaculatus (Schaller, 1783) (Coleoptera Cleridae Clerinae) EurasiaScience: Сollected Рapers, IX International Scientific-Practical in Russia. REDIA 101: 143–146; https://doi.org/10.19263/ Conference. Aktualnost.RF Publ., Moscow, 332pp [in Russian]. REDIA-101.18.19 Schnee, H. (2011). Beitrag zur Kenntnis der Biologie von Pseudogonalos Ruchin, A.B. & L.V. Egorov (2018b). Fauna of longicorn beetles hahnii (Spinola) (Hymenoptera, und Ichneumonidae). (Coleoptera: Cerambycidae) of Mordovia. Russian Entomol. Journal Entomologische Nachrichten und Berichte 55(1): 27–32. 27(2): 161–177; https://doi.org/10.15298/rusentj.27.2.07 Steiner, A.L. (1979). Digger wasp predatory behavior (Hymenoptera, Ruchin, A.B. & L.V. Egorov (2018c). Leptura aurulenta (Coleoptera, Sphecidae): hunting and capture by Oxybelus uniglumis Cerambycidae), a new record of a very rare species in Russia. Nature (Crabroninae: Oxybelini); a case of extremely concentrated stinging Conservation Research 3(1): 88–91; https://doi.org/10.24189/ pattern and prey nervous system. Canadian Journal of Zoology ncr.2018.003 57(5): 953–962; https://doi.org/10.1139/z79-120 Ruchin, A.B. & L.V. Egorov (2018d). On distribution of Mimela Straka, J. (2007). Vespoidea: Pompilidae (hrabalkovití). Acta holosericea (Fabricius, 1787) (Insecta, Scarabaeoidea, Entomologica Musei Nationalis Pragae Supplement 11: 111–131. Scarabaeidae, Rutelinae) in Russia and adjacent territories. Journal Tomaszewska, W., L.V. Egorov, A.B. Ruchin & D.V. Vlasov (2018). of Entomological and Acarological Research 50: 7390; https://doi. First record of the genus Clemmus (Coleoptera: Coccinelloidea, org/10.4081/jear.2018.7390 Anamorphidae) for the fauna of Russia. Nature Conservation Ruchin, A.B., L.V. Egorov & G.B. Semishin (2018). Fauna of click Research 3(3): 103–105; https://doi.org/10.24189/ncr.2018.016 beetles (Coleoptera: Elateridae) in the interfluve of rivers Moksha Vargot, E.V. (2016). Mordovia State Nature Reserve’s 80th anniversary. and Sura, Republic of Mordovia, Russia. Biodiversitas 19(4): 1352– Nature Conservation Research 1(2): 96–102; https://doi. 1365; https://doi.org/10.13057/biodiv/d190423 org/10.24189/ncr.2016.020 [in Russian] Ruchin, A.B., Egorov, L.V. & Semishin G.B. (2019). Ladybird beetles Yildirim, E., T. Ljubomirov, H. Özbek & M. Yüksel (2016). New data fauna (Coleoptera: Coccinellidae) of the Republic of Mordovia, on Spheciformes fauna (Hymenoptera: Ampulicidae, Sphecidae, Russia. Biodiversitas 20(2): 316-327; https://doi.org/10.13057/ Crabronidae) of Turkey. Journal of Insect Biodiversity 4(3): 1–51; biodiv/d200203 https://doi.org/10.12976/jib/2016.4.3

Threatened Taxa

13250 Journal of Threatened Taxa | www.threatenedtaxa.org | 15 February 2019 | 11(2): 13195–13250 Dr. Okan Külköylüoğlu, Abant Izzet Baysal University, Bolu, Turkey (Crustacea) Dr. Oguz Turkozan, Adnan Menderes University, Aydın, Turkey Dr. Jesse Leland, Southern Cross University, New South Wales, Australia (Crustacea) Dr. George Mathew, Kerala Forest Research Institute, Peechi, India Birds Dr. Mohilal Meitei, Manipur University, Camchipur, Manipur, India Dr. John C. Morse, Clemson University, Long Hall, Clemson, USA Dr. Hem Sagar Baral, Charles Sturt University, NSW Australia Late Dr. T.C. Narendran, (Retired) Professor, University of Calicut, Kerala, India Dr. Chris Bowden, Royal Society for the Protection of Birds, Sandy, UK Dr. John Noyes, Natural History Museum, London, UK Dr. Priya Davidar, Pondicherry University, Kalapet, Puducherry, India Dr. Albert G. Orr, Griffith University, Nathan, Australia (Odonata) Dr. J.W. Duckworth, IUCN SSC, Bath, UK Dr. Renkang Peng, Charles Darwin University, Darwin, Australia (Heteroptera) Dr. Rajah Jayapal, SACON, Coimbatore, Tamil Nadu, India Dr. Nancy van der Poorten, Toronto, Canada Dr. Rajiv S. Kalsi, M.L.N. College, Yamuna Nagar, Haryana, India Dr. C. Raghunathan, Zoological Survey of India, Andaman and Nicobar Islands Dr. Crawford Prentice, Nature Management Services, Jalan, Malaysia Dr. R. Ramanibai, Guindy Campus, Chennai, Tamil Nadu, India Dr. V. Santharam, Rishi Valley Education Centre, Chittoor Dt., Andhra Pradesh, India Dr. Brett C. Ratcliffe, University of Nebraska, Lincoln, USA Dr. C. Srinivasulu, Osmania University, Hyderabad, India Dr. Klaus Ruetzler, Smithsonian Institution, Washington, DC Dr. K.S. Gopi Sundar, International Crane Foundation, Baraboo, USA Dr. Kareen Schnabel, NIWA, Wellington, New Zealand (Crustacea) Dr. Gombobaatar Sundev, Professor of Ornithology, Ulaanbaatar, Mongolia Dr. R.M. Sharma, (Retd.) Scientist, Zoological Survey of India, Pune, India (Lepidoptera, Prof. Reuven Yosef, International Birding & Research Centre, Eilat, Israel Coleoptera) Dr. Taej Mundkur, Wetlands International, Wageningen, The Netherlands Dr. Peter Smetacek, Butterfly Research Centre, Bhimtal, India (Lepidoptera) Dr. Carol Inskipp, Bishop Auckland Co., Durham, UK Dr. Manju Siliwal, WILD, Coimbatore, Tamil Nadu, India (Araneae) Dr. Michael Hutchins, American Bird Conservancy, Washington, USA. Dr. G.P. Sinha, Botanical Survey of India, Allahabad, India (Lichens) Dr. V. Gokula, National College, Tiruchirappalli, Tamil Nadu, India Dr. K.G. Sivaramakrishnan, Madras Christian College, Chennai, Tamil Nadu, India Dr. K.A. Subramanian, Zoological Survey of India, New Alipore, Kolkata, India Mammals Dr. P.M. Sureshan, Zoological Survey of India, Kozhikode, Kerala, India (Hymenoptera) Dr. Martin B.D. Stiewe, The Natural History Museum, UK (Mantodea) Dr. Giovanni Amori, CNR - Institute of Ecosystem Studies, Rome, Italy Dr. R. Varatharajan, Manipur University, Imphal, Manipur, India Dr. Daniel Brito, Federal University of Goiás, Goiânia, Brazil Dr. Eduard Vives, Museu de Ciències Naturals de Barcelona, Terrassa, Dr. Anwaruddin Chowdhury, Guwahati, India Dr. John Veron, Coral Reef Foundation, Townsville, Australia Dr. P.S. Easa, Kerala Forest Research Institute, Peechi, India Dr. Hui Xiao, Chinese Academy of Sciences, Chaoyang, China Dr. Colin Groves, Australian National University, Canberra, Australia Dr. James Young, Hong Kong Lepidopterists’ Society, Hong Kong Dr. Cecília Kierulff, Victorville, California Dr. R. Sundararaj, Institute of Wood Science & Technology, Bengaluru, India (Isoptera) Dr. Kristin Leus, Copenhagen Zoo, Annuntiatenstraat, Merksem, Belgium Dr. M. Nithyanandan, Environmental Department, La Ala Al Kuwait Real Estate. Co. K.S.C., Dr. David Mallon, Zoological Society of London, UK Kuwait Dr. Antonio A. Mignucci-Giannoni, Universidad Interamericana de Puerto Rico, Puerto Dr. George O. Poinar, Oregon State University, Corvallis, USA Rico Dr. S. Arularasan, Annamalai University, Parangipettai, India (Molluscs) Dr. Sanjay Molur, WILD/ZOO, Coimbatore, India Dr. Himender Bharti, Punjabi University, Punjab, India (Hemiptera) Dr. Shomita Mukherjee, SACON, Coimbatore, Tamil Nadu, India Mr. Purnendu Roy, London, UK (Lepidoptera) Dr. P.O. Nameer, Kerala Agricultural University, Thrissur, Kerala, India Dr. Saito Motoki, The Butterfly Society of Japan, Tokyo, Japan Dr. Jill Pruetz, Iowa State University, Ames, USA Dr. Sanjay Sondhi, TITLI TRUST, Kalpavriksh, Dehradun, India (Lepidoptera) Dr. Ian Redmond, UNEP Convention on Migratory Species, Lansdown, UK Dr. Nguyen Thi Phuong Lien, Vietnam Academy of Science and Technology, Hanoi, Vietnam Dr. Heidi S. Riddle, Riddle’s Elephant and Wildlife Sanctuary, Arkansas, USA (Hymenoptera) Dr. Kumaran Sathasivam, Marine Mammal Conservation Network of India, India Dr. Xiaoli Tong, South China Agricultural University, Guangzhou, China (Thysanoptera) Dr. Karin Schwartz, George Mason University, Fairfax, Virginia. Dr. Nitin Kulkarni, Tropical Research Institute, Jabalpur, India (Orthoptera) Dr. Christoph Schwitzer, University of the West of England, Clifton, Bristol, BS8 3HA Dr. Robin Wen Jiang Ngiam, National Parks Board, Singapore (Odonata) Dr. Jodi L. Sedlock, Lawrence University, Appleton, USA Dr. Lional Monod, Natural History Museum of Geneva, Genève, Switzerland. Dr. Lala A.K. Singh, Bhubaneswar, Orissa, India Dr. Asheesh Shivam, Nehru Gram Bharti University, Allahabad, India Dr. Mewa Singh, Mysore University, Mysore, India Dr. Rosana Moreira da Rocha, Universidade Federal do Paraná, Curitiba, Brasil Dr. Meena Venkataraman, Mumbai, India Dr. Kurt R. Arnold, North Dakota State University, Saxony, Germany (Hemiptera) Dr. Erin Wessling, Max Planck Institute for Evolutionary Anthropology, Germany Dr. James M. Carpenter, American Museum of Natural History, New York, USA Dr. Dietmar Zinner, German Primate Center, Göttingen, Germany (Hymenoptera: Dr. A.J.T. Johnsingh, Nature Conservation Foundation, Mysuru and WWF-India, India Dr. David M. Claborn, Missouri State University, Springfield, USA (Diptera) Dr. Paul Racey, University of Exeter, Devon, UK Dr. Kareen Schnabel, Marine Biologist, Wellington, New Zealand Dr. Honnavalli N. Kumara, SACON, Anaikatty P.O., Coimbatore, Tamil Nadu, India Dr. Amazonas Chagas Júnior, Universidade Federal de Mato Grosso, Cuiabá, Brasil Dr. Ashwin Naidu, University of Arizona, Tucson, USA Dr. Marc W. Holderied, University of Bristol, Bristol, UK Fishes Dr. H. Raghuram, The American College, Madurai, Tamil Nadu, India Dr. David Olson, Zoological Society of London, UK Dr. Neelesh Dahanukar, IISER, Pune, Maharashtra, India Dr. Paul Bates, Harison Institute, Kent, UK Dr. Carl Ferraris, Smithsonian Institution, Portland, USA Dr. Hector Barrios-Garrido, James Cook University, Townsville, Australia Dr. M. Afzal Khan, Department of Zoology, Aligarh Muslim University, Aligarh, India Dr. Jim Sanderson, Small Wild Cat Conservation Foundation, Hartford, USA Dr. Topiltzin Contreras MacBeath, Universidad Autónoma del estado de Morelos, México Dr. Dan Challender, University of Kent, Canterbury, UK Dr. Heok Hee Ng, National University of Singapore, Science Drive, Singapore Dr. David Mallon, Manchester Metropolitan University, Derbyshire, UK Dr. Rajeev Raghavan, St. Albert’s College, Kochi, Kerala, India Dr. Lukas Rüber, Department of Vertebrates, Natural History Museum, Switzerland Other Disciplines Dr. Anjana Silva, Rajarata University of Sri Lanka, Saliyapura, Sri Lanka Dr. Robert D. Sluka, Chiltern Gateway Project, A Rocha UK, Southall, Middlesex, UK Dr. Aniruddha Belsare, Columbia MO 65203, USA (Veterinary) Dr. Kevin Smith, IUCN, Cambridge, UK Dr. S. Jayakumar, Pondicherry University, Puducherry, India (Climate Change) Dr. E. Vivekanandan, Central Marine Fisheries Research Institute, Chennai, India Dr. Jeff McNeely, IUCN, Gland, Switzerland (Communities) Dr. W. Vishwanath, Manipur University, Imphal, India Dr. Stephen D. Nash, Scientific Illustrator, State University of New York, NY, USA (Scientific Dr. J. Jerald Wilson, King Abdulaziz University, Jeddah, Saudi Arabia Illustrator) Dr. Davor Zanella, University of Zagreb, Zagreb, Dr. Mandar S. Paingankar, University of Pune, Pune, Maharashtra, India (Molecular) Dr. A. Biju Kumar, University of Kerala, Thiruvananthapuram, Kerala, India Dr. Jack Tordoff, Critical Ecosystem Partnership Fund, Arlington, USA (Communities) Dr. Frederic H. Martini, University of Hawaii at Manoa, Hanolulu, Hawaii Dr. Ulrike Streicher, University of Oregon, Eugene, USA (Veterinary) Dr. Hari Balasubramanian, EcoAdvisors, Nova Scotia, Canada (Communities) Amphibians

Dr. Indraneil Das, Sarawak, Malaysia Reviewers 2016–2018 Dr. Sushil K. Dutta, Indian Institute of Science, Bengaluru, Karnataka, India Due to pausity of space, the list of reviewers for 2016–2018 is available online. Dr. Annemarie Ohler, Muséum national d’Histoire naturelle, Paris, France

Reptiles

Late Dr. S. Bhupathy, SACON, Coimbatore, Tamil Nadu, India Dr. Llewellyn D. Densmore, Texas Tech University, Lubbock, USA Dr. Eric Smith, University of Texas, Arlington, USA Dr. Gernot Vogel, Heidelberg, Germany Dr. Anders G.J. Rhodin, Chelonian Research Foundation, Lunenburg, USA Dr. Raju Vyas, Vadodara, Gujarat, India Dr. Pritpal S. Soorae, Environment Agency, Abu Dubai, UAE. Dr. Olivier S.G. Pauwels, Royal Belgian Institute of Natural Sciences, Brussels, Belgium The opinions expressed by the authors do not reflect the views of the Dr. Anders G.J. Rhodin, Chelonian Research Foundation, Lunenburg, USA Journal of Threatened Taxa, Wildlife Information Liaison Development Society, Zoo Outreach Organization, or any of the partners. The journal, the publisher, the host, and the partners are not responsible for the accuracy of the political Journal of Threatened Taxa is indexed/abstracted in Bibliography boundaries shown in the maps by the authors. of Systematic Mycology, Biological Abstracts, BIOSIS Previews, CAB Abstracts, EBSCO, Google Scholar, Index Copernicus, Index Fungorum, JournalSeek, National Academy of Agricultural Sciences, NewJour, OCLC Print copies of the Journal are available at cost. Write to: WorldCat, SCOPUS, Stanford University Libraries, Virtual Library of Biol- The Managing Editor, JoTT, c/o Wildlife Information Liaison Development Society, ogy, Zoological Records. No. 12, Thiruvannamalai Nagar, Saravanampatti - Kalapatti Road, Saravanampatti, Coimbatore, Tamil Nadu 641035, India NAAS rating (India) 5.10 [email protected] PLATINUM The Journal of Threatened Taxa (JoTT) is dedicated to building evidence for conservation globally by publishing peer-reviewed articles online every month at a reasonably rapid rate at www.threatenedtaxa.org. OPEN ACCESS All articles published in JoTT are registered under Creative Commons Attribution 4.0 International License unless otherwise mentioned. JoTT allows allows unrestricted use, reproduction, and distribution of articles in any medium by providing adequate credit to the author(s) and the source of publication.

ISSN 0974-7907 (Online) | ISSN 0974-7893 (Print)

February 2019 | Vol. 11 | No. 2 | Pages: 13195–13250 Date of Publication: 15 February 2019 (Online & Print) www.threatenedtaxa.org DOI: 10.11609/jott.2019.11.2.13195-13250

Monograph

Wasp fauna (Hymenoptera: Bethylidae, Chrysididae, Dryinidae, Tiphiidae, Mutillidae, Scoliidae, Pompilidae, Vespidae, Sphecidae, Crabronidae & Trigonalyidae) of Mordovia State Nature Reserve and its surroundings in Russia – Аlexander Ruchin & Аlexander Antropov, Pp. 13195–13250

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Threatened Taxa