Pseudepipona herrichii (de Saussure, 1856)

Fateryga, Alexander V., 2022, Revision of the Pseudepipona herrichii-group of the eumenine wasps (Hymenoptera: Vespidae: Eumeninae) with the description of two new species from China and Russia, Zootaxa 5154 (2), pp. 101-126 : 110-114

Pseudepipona herrichii (de Saussure, 1856)

( Figs 10–12 , 26 , 29–38 )

Odynerus variegatus Herrich-Schäffer, 1839: 16–17 , ♀; type locality: “auf Reseda lutea bei Regensburg” [ Germany]; nom. praeocc., nec Odynerus variegatus ( Fabricius, 1793) .

Odynerus herrichii de Saussure, 1856: 309 , replacement name for O. variegatus Herrich-Schäffer, 1839 , nec O. variegatus ( Fabricius, 1793) .

Odynerus basalis Smith, 1857: 58 , ♀; type locality: “Polish Ukraine ”; synonymized by E. Saunders in Mortimer, 1908: 236.

Pseudepipona herrichii var. derufata Blüthgen, 1951: 184–186 , ♀ ♂; type locality: “ Eriwan ” [ Armenia]; synonymized by van der Vecht & Fischer, 1972: 83–84.

Pseudepipona variegata: Blüthgen, 1952: 29 .

Pseudepipona herrichii: Giordani Soika, 1970: 130 .

Pseudepipona herrichii afromontana Gusenleitner, 1977: 142–143 , ♀ ♂; type locality: “Oukaimeden, 2600–2800 m ” [Morocco], syn. nov.

Material examined. MOROCCO: H. Atlas, Oukaïmeden , 2700–2900 m, 12.VII.1975, 1 ♀, leg. J. Gusenleitner [ FSCV] (paratype of Pseudepipona herrichii afromontana ); ibid., 12.VII.1975, 1 ♂, leg. J. Gusenleitner [ OLML] (paratype of P. herrichii afromontana ) . ALGERIA: Algier, 1 ♀ [ ZISP] . SPAIN: Avila, 1865, 1 ♂, leg. G. Sedlitz [ ZISP]; Teruel , Albarracín: Vallejo Largo, 1170 m, 23. VI.1996, 1 ♂, leg. L. Castro [ CAFK]; Teruel , Camarena: Belneario, 1350 m, 22. VI.1996, 1 ♂, leg. L. Castro [ CAFK]; Teruel, Nacimiento del r. Tajo: Nava Seca, 1585 m, 30. VI.1990, 1 ♀, leg. L. Castro [ CAFK]; ibid., 10.VII.1996, 1 ♂, leg. L. Castro [ CAFK] . SLOVAKIA: Čenkov u Štúrova, 28.VII.1955, 1 ♂, leg. Bouček [ OLML]; Somotor , 5.VII.1952, 1 ♀ [ OLML] . MOLDOVA: Chișinău, 23. V.1899, 1 ♂, leg. Zubovskiy [ ZISP]. UKRAINE: Kuryany , 19.VII.1886, 1 ♀ [ ZISP]; Poltava Governement, Krasnograd, 7. V.1923, 1 ♂, leg. F. Lukyanovich [ ZISP]; Vicinity of Kiev, Kruglik, 3. VI .1979, 1 ♀, leg. A.S. Lelej [ FSCV]; ibid., 12. VI.1979, 1 ♀, leg. A.S. Lelej [ FSCV] . RUSSIA. Kirov Province: Malmuzh, 15–20. V .1948, 1 ♂ [ ZISP]. Belgorod Province: Borisovka, 28. VI .1958, 1 ♂ [ ZISP]. Saratov Province: Samara Governement, Nikolaevsk Uezd , 2 ♀, 1 ♂, leg. V . Bostanzhoglo [ ZISP]; ibid., 6. VI , 5 ♂, leg. V . Bostanzhoglo [ ZISP]; ibid., 7. VI , 1 ♀, 1 ♂, leg. V . Bostanzhoglo [ ZISP]; ibid., 16. VI , 1 ♀, leg. V . Bostanzhoglo [ ZISP]; ibid., 17. VI , 1 ♀, leg. V . Bostanzhoglo [ ZISP]; ibid., 18. VI , 3 ♀, 2 ♂, leg. V . Bostanzhoglo [ ZISP]; ibid., 23. VI , 2 ♀, 1 ♂, leg. V . Bostanzhoglo [ ZISP]; ibid., 1.VII, 2 ♀, 1 ♂, leg. V . Bostanzhoglo [ ZISP]; ibid., 6.VII, 1 ♀, leg. V . Bostanzhoglo [ ZISP]; ibid., 7.VII, 1 ♀, leg. V. Bostanzhoglo [ ZISP]; Samara Governement , Nikolayevsk Uezd, Roslyakovo, 24. VI.1898, 1 ♂, leg. E. Klements [ ZISP]. Volgograd Province: Uryupinskaya [currently Uryupinsk], Holy Mountains , 12. VI.1910, 1 ♀, leg. N.A. Popov [ ZISP]. Astrakhan Province: Astrachan [= Astrakhan], 1 ♂ [ ZISP]; Bogdo , 3.VII.1877, 1 ♂ [ ZISP]. Stavropol Territory: Pyatigorsk, 1886, 1 ♀ [ ZISP]. Ingushetia: Salgi, 5462′, 29.VII.1927, 1 ♂, leg. A. Kiritchenko [ ZISP]. Dagestan: Petrovsk [currently Makhachkala], 23. V .1901, 1 ♂, leg. Beckmann [ ZISP]; ibid., 24. V .1901, 2 ♀, 1 ♂, leg. Beckmann [ ZISP]; ibid., 25. V .1901, 2 ♂, leg. Beckmann [ ZISP]; ibid., 27. V .1901, 1 ♀, 1 ♂, leg. Beckmann [ ZISP]; Crimea: Alma-Thal, 1 ♀ [ ZISP]; Feodosiya , Lisya Bay , 11. VI .2007, 3 ♀, leg. A. V . Fateryga [ CFUS]; ibid., 13. VI .2007, 5 ♀, leg. A. V . Fateryga [ CFUS]; ibid., 13. VI .2007, 1 ♀, leg. A. V . Fateryga [ CAFK]; ibid., 16. VI .2007, 2 ♀, leg. A. V . Fateryga [ CFUS]; ibid., 26. VI .2009, 2 ♀, leg. A. V . Fateryga [ CFUS]; ibid., 12. VI .2010, 2 ♀, leg. A. V . Fateryga [ CFUS]; ibid., 16. VI .2011, 1 ♀, leg. A. V . Fateryga [ CFUS]; Gromovka, 6– 8. VI .1996, 2 ♂, leg. S.P. Ivanov [ CFUS]; Kerch Peninsula , Cape Chauda, on Butomus umbellatus , 9. VI.2005, 1 ♀, leg. S.P. Ivanov [ CFUS]; Koyash [Lake], 23.VII.1900, 1 ♂ [ ZISP]; Nizhnigorskiy District, Peny , 12.IX.1978, 1 ♀, leg. S.A. Mosyakin [ SMS]; Sevastopol , Kazachya Bay , 7.VII.2005, 1 ♀, leg. I.S. Turbanov [ CFUS]; Sevastopol, Khersones, 27. VI .1912, 8 ♀, leg. V . Pliginskiy [ ZISP]; ibid., 29. VI .1912, 4 ♀, leg. V . Pliginskiy [ ZISP]; ibid., 30. VI .1912, 1 ♀, leg. V . Pliginskiy [ ZISP]; ibid., 21. VI .1914, 1 ♀, leg. V . Pliginskiy [ ZISP]; Simferopol , 18.VII.1900, 1 ♀ [ ZISP]; ibid., 15. V .1904, 1 ♂ [ ZISP]; ibid., 15. VI .1928, 1 ♀, 1 ♂, leg. A.I. Kazanskiy [ ZISP]; ibid., 20.VII.2002, 1 ♀, leg. A. V . Fateryga [ CAFK]; ibid., 20.VII.2002, 3 ♀, leg. A. V . Fateryga [ CFUS]; ibid., 21.VII.2002, 4 ♀, leg. A. V . Fateryga [ CFUS]; Simferopol , Petrovskaya Balka, 13. VI .2003, 1 ♀, leg. A. V . Fateryga [ CAFK]; ibid., 13. VI .2003, 3 ♀, leg. A. V . Fateryga [ CFUS]; Simferopol, Salgir River valley, 13.VII.1952, 1 ♂ [ OSZ]; Sudak , 7.VII.2005, 1 ♀, leg. A. V . Fateryga [ CFUS]; Tarkhankut Peninsula, 24. VI .2003, 11 ♀, leg. A. V . Fateryga [ CFUS]; Tarkhankut Peninsula, Bolshoy Kastel , 4.VII.2014, 1 ♀, leg. V . Yu. Zhidkov [ CFUS]; ibid., 22.VII.2015, 1 ♀, leg. A. V . Fateryga [ CFUS]; Tarkhankut Peninsula , Kipchak, 24. VI .2005, 2 ♀, leg. A. V . Fateryga [ CFUS]; ibid., 26. VI .2005, 7 ♀, leg. D. V . Puzanov [ DPS]; ibid., 21. VI .2007, 1 ♀, leg. A. V . Fateryga [ CFUS]; ibid., 22. VI .2007, 7 ♀, leg. A. V . Fateryga [ CFUS]; ibid., 22. VI .2007, 1 ♀, leg. A. V . Fateryga [ CAFK]; ibid., 25. VI .2007, 1 ♀, leg. D. V . Puzanov [ CAFK] (with reddish spots on T2 ); ibid., 18. VI.2008, 1 ♂, leg. A. V. Fateryga [ CAFK]; ibid., 18. VI.2008, 1 ♀, leg. A. V. Fateryga [ CFUS]; ibid., 19. VI.2008, 2 ♂, leg. A. V. Fateryga [ CAFK]; ibid., 20. VI.2008, 6 ♀, leg. A. V. Fateryga [ CFUS]; ibid., 21. VI.2008, 2 ♂, leg. A. V. Fateryga [ CAFK]; ibid., 22. VI.2008, 4 ♀, 1 ♂, leg. A. V. Fateryga [ CFUS]; ibid., 22. VI.2008, 11 ♀, leg. D. V. Puzanov [ DPS]; Vicinity of Simferopol , 2. VI.1928, 1 ♂ [ OSZ]; ibid., VII–VIII.1898, 1 ♀, leg. Graftio [ ZISP]; Western Coast, Cape Lukull , 8.VII.2015, 1 ♀, leg. A. V. Fateryga [ CFUS]. Bashkortostan: Ufa Governement, Belebey Uezd, 2 ♀, leg. B. Kazanskiy [ ZISP]. Orenburg Province: Upper Dneprovka , 2.VII.1934, 1 ♂, leg. L. Zimin [ ZISP]; Vicinity of Orenburg, 10.VII.1924, 1 ♀, leg. P.A. Vorontsovskiy [ ZISP]; ibid., 2. VI.1927, 1 ♂, leg. P.A. Vorontsovskiy [ ZISP]; ibid., 24. VI.1927, 1 ♀, leg. P.A. Vorontsovskiy [ ZISP]; ibid., 13.VIII.1932, 4 ♀, leg. P.A. Vorontsovskiy [ ZISP]. Omsk Province: Omsk, 10–22. VI.1926, 1 ♂, leg. Bey-Bienko [ ZISP] . GEORGIA: Lagodechi , 1 ♀, 1 ♂ [ ZISP]. ARMENIA: Eriwan [= Yerevan], 1 ♂ [ ZISP]. AZERBAIJAN: Adshikent [= Hacıkənd], 1 ♂ [ ZISP]; Nakhchivan AR, Babek , Goynuk, 39°18′N, 45°40′E, 1680 m, 12. VI.2019, 1 ♂, leg. M.Yu. Proshchalykin, Kh.A. Aliyev & M.M. Maharramov [ CAFK] ( Fateryga et al. 2021, corresponds to var. derufata ); Ordubad ( Arax ), 1892, 1 ♂, leg. Reitter [ ZISP] (corresponds to var. derufata ). UZBEKISTAN : Aman-Kutan , 22.VII.1931, 1 ♂, leg. V . Gussakovskij [ ZISP]; Chatkalskiy Range 30 km N Parkent, Aksagata Canyon, 7–10. VI .1982, 1 ♂, leg. V . L. Kazenas [ FSCV] . KYRGYZSTAN: Issyk-Kul Lake , Dolinka [currently Kara-Oy], 7. VI .1973, 1 ♂, leg. Yu. Tarbinskiy [ FSCV]; Issyk-Kul Lake, N coast, vicinity of Bosteri , 23.VII.1983, 1 ♂, leg. N. V . Kurzenko [ FSCV]; S slopes of Moldo-Tau near Shavyr Landmark , 18.VII.1971, 1 ♀, leg. Yu. Tarbinskiy [ FSCV]; Semirechye [= Zhetysu], Pishpekskiy Uezd, Uchimchik, 11.VIII.1913, 1 ♀, leg. V. Chernavin [ ZISP]; Vicinity of Kazarman, 2000 m, 13.VII.1972, 3 ♀, leg. Yu. Tarbinskiy [ FSCV]; Vicinity of Tasharyk , 15 km E Talas, 3.VIII.1983, 1 ♀, leg. V.L. Kazenas [ FSCV] . KAZAKHSTAN. Akmola Province: Akmolinsk [currently Nur-Sultan], 20. VI.1931, 1 ♂, leg. V. B. Popov [ ZISP]; Bayzhanzhal, 7.VII.1957, 1 ♀, leg. Rudolf [ ZISP]; Kokshetau, near Tersakan River , 28. V.1957, 1 ♂, leg. Rudolf [ ZISP]; ibid., 1.VII.1957, 1 ♀, leg. Rudolf [ ZISP]; ibid., 29.VII.1958, 1 ♀, leg. Rudolf [ ZISP]; Tersakan River, 10 km from Kokshetau, 9.VII.1957, 1 ♀, leg. Rudolf [ ZISP]; Vicinity o Kiyma , 7.VII.1977, 1 ♀, 1 ♂, leg. V . L. Kazenas [ FSCV]; Vicinity of Derzhavinsk , 4.VII.1977, 1 ♂, leg. V . L. Kazenas [ FSCV]. Aktobe Province: Ber Tschogur, Mugodjargebirge [currently Birshoghyr], 1 ♀, leg. L. Wollmann [ ZISP]. Almaty Province: 20 km W Alma-Ata, 26. VI.1985, 1 ♀, leg. V. L. Kazenas [ FSCV]; Ketmen Landmark, north slopes near Podgornoye [currently Kyrgyzsay], 22.VII.1967, 1 ♀, leg. P.A. Lehr [ FSCV]; Ketmen-Tau, Bolshoy Kirgiz-Say Landmark, N slopes, 15.VII.1968, 1 ♂ [ FSCV]; N foothills of Zhetykol Range, 29.VII.1971, 1 ♀, leg. N. V. Kurzenko [ FSCV]; Sarkand [District], Topolevka, Dzhungarian Alatau, 15. VI.957, 1 ♂, leg. Kerzhner [ ZISP]; Trans-Ili Alatau , Bolshoy Dalan Canyon, 2. VI.1971, 1 ♂, leg. N. V. Kurzenko [ FSCV]; ibid., 19. VI .1971, 1 ♀, 1 ♂, leg. N. V . Kurzenko [ FSCV]; Trans-Ili Alatau, vicinity of Alma-Ata , 6.VII.1974, 2 ♀, 20 ♂, leg. N. V . Kurzenko [ FSCV]; Vicinity of Alma-Ata, Aksay , 12.VII.1983, 1 ♀, 1 ♂, leg. N. V . Kurzenko [ FSCV]; Atyrau Province, Vicinity of Sagiz , 2.VII.1969, 1 ♂, leg. V . L. Kazenas [ FSCV]. East Kazakhstan Province: 10 km NW UstKamenogorsk , 2.VII.1979, 1 ♀, leg. V. L. Kazenas [ FSCV]; 18 km NE Ayaguz [= Ayagoz ], 1–8.VII.1986, 1 ♀, leg. V . L. Kazenas [ FSCV]; 50 km NE Ayaguz [= Ayagoz], 19. VI.1986, 1 ♂, leg. V. L. Kazenas [ FSCV]; 9–12 km W Ayaguz [= Ayagoz ], 19.VII.1986, 2 ♀, leg. V. L. Kazenas [ FSCV]; Chagan-Obo, Saur, 12–14.VII.1910, 1 ♂, leg. B. Karavayev [ ZISP]; Kazakh Uplands , Kaynar, 28. V .1974, 1 ♂, leg. N. V. Kurzenko [ FSCV]; Semipalatinsk [currently Semey], 2 ♂ [ ZISP]; Urdzhar, 29. VI.1962, 1 ♂, leg. V.L. Kazenas [ FSCV]. Jambyl Province: Aleksandrovskiy Range , Aulie-Ata Uezd, Kilan Canyon above Dmitriyevka, 16. VI.1913, 1 ♀, leg. V. Chernavin [ ZISP]. Karaganda Province: 55–60 km SEE Karkaralinsk , 21. VI .1978, 2 ♂, leg. V . L. Kazenas [ FSCV]; Kazakh Uplands , 0–200 km N Balkhash Lake, 25. V .1969, 1 ♀, leg. V . L. Kazenas [ FSCV]. Kostanay Province: Kunduzda River , inflow of Nura, 16. VI .1900, 1 ♀, leg. Balykleysk [iy] [ ZISP]; ibid., 22. VI –1.VII.1900, 1 ♀, leg. Balykleysk [iy] [ ZISP]; ibid., 29. VI –1.VII.1900, 1 ♀, leg. Balykleysk [iy] [ ZISP]; Vicinity of Arkalyk , 22. VI .1977, 1 ♂, leg. V . L. Kazenas [ FSCV]. North Kazakhstan Province: Omsk Uezd, W of Teke Lake, 21.VIII.1899, 1 ♀, leg. Ignatov [ ZISP]. COUNTRY UNCERTAIN: Caucasus , 1 ♂ [ ZISP] .

Observations on iNaturalist. RUSSIA. Crimea : Tarkhankut Peninsula, 45.49°N, 32.60°E, 13. VI.2014, 1 ♀ , A. V. Fateryga (https://www.inaturalist.org/observations/68380454); ibid., 20. VI.2008, 2 ♀ , A. V. Fateryga (https:// www.inaturalist.org/observations/68370002, https://www.inaturalist.org/observations/68691548); ibid., 6. VII.2018, 1 ♀ , A. V. Fateryga (https://www.inaturalist.org/observations/68691549) .

Additional literature records. MOROCCO: H. Atlas, Oukaïmeden, 2800–3000 m ( Gusenleitner 1977); H. Atlas, Oukaïmeden, 4 km E, 2300 m ( Gusenleitner 1977). GREAT BRITAIN: Between Wareham and Studland ( Spooner 1934); Bloxworth Heath ( Spooner 1934); In close proximity to Swanage, Studland Heath ( Spooner 1934); Isle of Purbek, Studland ( Spooner 1934); Isle of Purbek, Wareham ( Spooner 1934); Stoborough Heath, between Wareham and Corfe ( Spooner 1934). THE NETHERLANDS: Roermond ( Blüthgen 1961); Roozendaal ( Blüthgen 1961). FRANCE: Aveyron ( Berland 1928); Basses-Alpes: Escaffarels ( Berland 1928); Lyon ( Blüthgen 1961). SWITZERLAND: Wallis bei Sierre und Sion ( Blüthgen 1961). AUSTRIA: Guntramsdorfer Sandgruben ( Blüthgen 1961); Neusiedlersee ( Blüthgen 1961); Stammersdorf und Oberweiden ( Blüthgen 1961). PORTUGAL: Algarve, Aljezur, Ponta da Atalaia ( Baldock et al. 2020); Algarve, Aljezur, W of Vale de Telha ( Baldock et al. 2020); Algarve, Ponta da Atalia ( Baldock et al. 2020). SPAIN: Bayona und Montseny ( Blüthgen 1961); Cuenca ( Blüthgen 1951); Losilla bei Albarracin ( Blüthgen 1951); Montarco ( Blüthgen 1961). ITALY: Bei Bozen ( Blüthgen 1961); Calabrien ( Blüthgen 1961). SLOVENIA ( Gusenleitner 2013). CROATIA ( Gusenleitner 2013). MONTENEGRO ( Gusenleitner 2013). SERBIA ( Gusenleitner 2013). GREECE: Attica ( Blüthgen 1951); Syra ( Blüthgen 1951). HUNGARY: Budapest ( Móczár 1938); Isaszeg ( Móczár 1938); Péczel ( Móczár 1938). CZECH REPUBLIC ( Gusenleitner 2013). ROMANIA: Nagyvárad ( Móczár 1938). UKRAINE: Donetsk Province, Nozoazovsk District, Khomutovskaya Steppe Preserve ( Amolin 2009); Donetsk Province, Starobeshevo District, vicinity of Razdolnoye ( Amolin 2009); Lugansk Province, Sverdlovsk District, Provalskaya Steppe Preserve ( Amolin 2009); SE outskirts of Donetsk ( Amolin 2009). BELARUS: Provincia Mohileviensis ( Arnold 1902). RUSSIA. Rostov Province: Azov District, Ust-Koysug ( Amolin & Artokhin 2014); Rostov-on-Don, Botanical Garden ( Amolin & Artokhin 2014); Semikarakorsk District, Kirsanovka ( Amolin & Artokhin 2014); Sholokhovskiy District, Veshenskaya ( Amolin & Artokhin 2014). GEORGIA: Achalzik ( Blüthgen 1951); Lagodechi ( Blüthgen 1951). TURKEY: Ankara, Elma Dagi, 1000 m ( Giordani Soika 1970a); Ankara, Idris Dagi, 1300 m ( Giordani Soika 1970a); Bayburt: Kopdağı, Şehitler Çeşmesi, 2400 m ( Yildirim & Gusenleitner 2009); Erzurum, Kopdagi Gecidi, 6500 piedi ( Giordani Soika 1970a); Erzurum: Oltu, 1455 m ( Yildirim & Gusenleitner 2015); Hakkari: Yüksekova, Akçalı, 37°21′N, 44°03′E, 1700 m ( Yildirim & Gusenleitner 2012); Kares, Ararat, Karasu Spring, 2400 piedi ( Giordani Soika 1970a); Kars, Ilgaz Dagi, 1300 m ( Giordani Soika 1970a); Province Hakkari: Yüksekova—Semdinli, Sumpfwiese ( Madl 1997). UZBEKISTAN: Schachimardan ( Blüthgen 1951).

Diagnosis. Females of this species can be distinguished from other members of the P. herrichii -group by the following combination of characters: clypeus 1.10–1.15× as wide as long, black with a basal whitish or yellow band; scutellum with an apical band usually thicker than that on the metanotum and somewhat interrupted medially, sometimes the band on the metanotum or rarely both bands are reduced; propodeum with a weakly developed transverse carina between the shelf and concavity forming laterally rather right angles; propodeal valvula rounded posteriorly; T1 without three small longitudinal carinae. In the males, the clypeus is also 1.10–1.15× as wide as long; cuspis with a very long and relatively narrow dorsal process which is longer than its width at the base; base of the digitus attached somewhat below the top of the cuspis apex so that the latter is protruding as a short free acute spine; aedeagus with a median expansion; ventral lobe of the aedeagus small, in lateral view approximately as large as the median expansion.

Distribution. Morocco, Algeria, Great Britain, The Netherlands, France, Switzerland, Germany, Austria, Portugal, Spain, Italy, Slovenia, Croatia, Montenegro, Serbia, Greece, Hungary, Czech Republic, Slovakia, Romania, Moldova, Ukraine, Belarus, Russia (Kirov Province, Belgorod Province, Saratov Province, Rostov Province, Volgograd Province, Astrakhan Province, Stavropol Territory, Ingushetia, Dagestan, Crimea, Bashkortostan, Orenburg Province, Omsk Province), Georgia, Armenia, Azerbaijan, Turkey, Uzbekistan, Kyrgyzstan, Kazakhstan (Akmola Province, Aktobe Province, Almaty Province, East Kazakhstan Province, Jambyl Province, Karaganda Province, Kostanay Province, North Kazakhstan Province) ( Fig. 26 ). The reports from the Novosibirsk Province of Russia (e.g., Fateryga & Mokrousov 2019; Fateryga et al. 2020) probably refer to P. aldrichi , as are most records from Mongolia and China (e.g., Buyanjargal et al. 2017; Bai et al. 2021) except a few that could actually refer to P. valentinae sp. nov. The presence of the true P. herrichii in China is, however, still possible because it is distributed in the neighboring Kyrgyzstan and the eastern provinces of Kazakhstan. The reports of P. herrichii from Syria ( Fateryga & Mokrousov 2019; Fateryga et al. 2020; Rahmani et al. 2020) were typos (actually referring to Syra in Greece, Blüthgen 1951). The reports from Turkmenistan, in the same papers cited above, were not confirmed by the present study, however, its presence there is possible. The report from Iran ( Gusenleitner 1998) also needs confirmation (see Rahmani et al. 2020). All reports from North America refer to P. aldrichi (see above).

Remarks. This species is very polymorphic in coloration. Specimens from Morocco ( P. herrichii afromontana ), Portugal, Spain, and Azerbaijan ( P.herrichii derufata ) are black with yellow ( Figs 29, 37 ); those from Azerbaijan ( Fig. 37 and Fig. 27 in Fateryga et al. 2021) are, in addition, characterized by a coarser punctation of the metasomal terga. Typical specimens from the main part of the species range are black with whitish bands on the body and mostly reddish legs and T1 ( Figs 31, 33 ). However, several different color forms can be present even in the same region such as the Crimea (cf. Figs 31, 35, 36 ) or Tien Shan in Kyrgyzstan and the Almaty Province of Kazakhstan. The male genitalia are also somewhat variable in the size of the dorsal process of cuspis and the shape of the median expansion of the aedeagus (particularly, this expansion is more acute in the specimens from the westernmost parts of the species range) but these variations seem to be not correlated with the coloration.

Bionomics. Nests of this species were documented mostly in Great Britain by Mortimer (1908) and Spooner (1934). They noted that the nests were excavated close to each other in sandy soil. Females excavated their nests by extracting sand particles from the burrow and depositing them in approximately 1–2 inches (= 2.5–5.1 cm) from the entrance. The depth of the nests were about 1.5–2.5 inches (= 3.8–6.4 cm); there was no turret at the nest entrance. Apparently, the females used water to moisten the substrate during the excavation. In the Crimea, females were often observed collecting water ( Fig. 38 ), including water from the sea ( Fateryga & Zhidkov 2012). Spooner (1934) reported that the number of cells in the nests was from one to three; an illustrated three-celled nest had a branched or perhaps a linear-branched structure. The egg was suspended with a thread. Complete nests were sealed by filling the burrow with sand particles, apparently without their bonding with liquid. Baldock et al. (2020) published a photo of a female excavating its nest burrow in Portugal, also in sandy soil. Provision of P. herrichii is moth caterpillars, probably in the genus Depressaria Haworth ( Lepidoptera : Depressariidae ) ( Mortimer 1908). They live on the grass, in cocoon-like webs from which the wasp females have to capture them. Ferton (1909) reported a female searching for caterpillars on the ground surface under the grass but that case might refer to a female that had lost its prey and tried to find it (cf. Spooner 1934). The number of prey items recorded per cell was 8–13 ( Spooner 1934). Adult feeding of P. herrichii was recorded on flowers of Reseda lutea L. ( Resedaceae ) in Germany ( Herrich-Schäffer 1839) and Butomus umbellatus L. ( Butomaceae ) in the Crimea (see Material examined).

Figures

FIGURES 7–24. Male genitalia of Pseudepipona spp. 7–9, P. aldrichi (Fox, 1892); 10–12, P. herrichii (de Saussure, 1856); 13–15, P. kozhevnikovi (Kostylev, 1927); 16–18, P. tingjingliae Fateryga, sp. nov.; 19–21, P. tricarinata (Kokujev, 1913); 22–24, P. valentinae Fateryga, sp. nov. 7, 10, 13, 16, 19, 22, left paramere and volsella in lateral view from inside (bd, base of digitus; ca, cuspis apex; dp, dorsal process of cuspis); 8, 11, 14, 17, 20, 23, aedeagus in dorsal view (me, median expansion); 9, 12, 15, 18, 21, 24, aedeagus in lateral view (vl, ventral lobe). Scale bar 1 mm.

FIGURES 25–28. Distribution of Pseudepipona spp. 25, P. aldrichi (Fox, 1892); 26, P. herrichii (de Saussure, 1856); 27, P. kozhevnikovi (Kostylev, 1927); 28, P. tingjingliae Fateryga, sp. nov. (black dots), P. tricarinata (Kokujev, 1913) (white dots), and P. valentinae Fateryga, sp. nov. (grey dots).

FIGURES 29–37. Pseudepipona herrichii (de Saussure, 1856). 29, 31, 33, 35–37, habitus in dorsal view (29, 31, 35, 36, ♀; 33, 37, ♂); 30, 32, 34, head in frontal view (30, 32, ♀; 34, ♂). 29, 30, Morocco (paratype of P. herrichii afromontana Gusenleitner, 1977); 31–36, Russia: Crimea; 37, Azerbaijan. Scale bars 1 mm.

FIGURES 38–39. Females of Pseudepipona spp. 38, P. herrichii (de Saussure, 1856) from the Crimea (on water), photo by the author; 39, P. valentinae Fateryga, sp. nov. from Beijing (with a prey), photo by Home Mountain from the iNaturalist web-site (https://www.inaturalist.org/observations/102650425), distributed under the terms of the Creative Commons BY-NC 4.0 license (https://creativecommons.org/licenses/by-nc/4.0/).