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Order Hymenoptera. In: Zhang, Z.-Q. (Ed.) Animal Biodiversity: An Outline of Higher-level Classification and Survey of Taxonomic Richness (Addenda 2013)



An updated classification of the order Hymenoptera is provided with the current numbers of genera and species described so far specified. The order is composed of 2 suborders, 27 superfamilies, 132 families, 8423 extant genera with an additional 685 extinct genera. Considered one of the most species-rich insects orders a total of 153088 extant species have been described, in addition to 2429 extinct species.
Accepted by Zhi-Qiang Zhang: 18 Aug. 2013; published: 30 Aug. 2013
Licensed under a Creative Commons Attribution License
ISSN 1175-5326 (print edition)
1175-5334 (online edition)
Copyright © 2013 Magnolia Press
Zootaxa 3703 (1): 051062
zoota xa
Order Hymenoptera*
Universidade Federal do Espírito Santo, Depto. de Ciências Biológicas, Av. Fernando Ferrari, s/n – Goiabeiras, 29075-010 Vitoria,
ES, BRAZIL, e-mail:
Department of Entomology, Pennsylvania State University, 501 ASI Building, University Park, PA 16802, USA,
Division of Entomology, Natural History Museum, 1501 Crestline Drive – Suite 140, University of Kansas, Lawrence, KS 66045,
Swedish Museum of Natural History, Department of Entomology, Box 50007, SE-104 05 Stockholm, SWEDEN,
Canadian National Collection of Insects (CNC), Agriculture and Agri-Food Canada, 960 Carling Avenue, Ottawa, ON, K1A 0C6,
School of Earth & Environmental Sciences, G15d Darling Building, The University of Adelaide, 5005, AUSTRALIA,
Department of Evolution, Ecology & Organismal Biology, Director C.A. Triplehorn Insect Collection, The Ohio State University,
1315 Kinnear Road, Columbus, OH 43212, USA,
Institute of Biology and Soil Science, Russian Academy of Sciences, Vladivostok-22, 690022, RUSSIA, e-mail:
Department of Biology, 257 S 1400 E, University of Utah, Salt Lake City, UT 84112, USA, e-mail:
Museum für Naturkunde, Leibniz-Institut für Evolutions- und Biodiversitätsforschung, Invalidenstraße 43, 10115 Berlin, GERMANY,
current address Übersee-Museum Bremen, Stiftung öffentlichen Rechts, Bahnhofsplatz 13, 28195 Bremen, GERMANY,
Department of Entomology, Pennsylvania State University, 501 ASI Building, University Park, PA 16802, USA,
Museum für Naturkunde, Leibniz-Institut für Evolutions- und Biodiversitätsforschung, Invalidenstraße 43, 10115 Berlin, GERMANY,
Department of Bioscience, Aarhus University, Ny Munkegade 114, Bldg. 1540, DK-8000 Aarhus C, DENMARK,
Senckenberg Deutsches Entomologisches Institut (SDEI), Eberswalder Str. 90, 15374 Müncheberg, GERMANY,
403-1220 Merivale Road, Ottawa, Ontario K1Z 8P2, CANADA, e-mail:
** Corresponding and compiling author.
* In: Zhang, Z.-Q. (Ed.) Animal Biodiversity: An Outline of Higher-level Classification and Survey of
Taxonomic Richness (Addenda 2013). Zootaxa, 3703, 1–82.
An updated classification of the order Hymenoptera is provided with the current numbers of genera and species
described so far specified. The order is composed of 2 suborders, 27 superfamilies, 132 families, 8423 extant genera with
an additional 685 extinct genera. Considered one of the most species-rich insects orders a total of 153088 extant species
have been described, in addition to 2429 extinct species.
Key words: Hymenoptera, wasps, bees, ants, classification, diversity
52 · Zootaxa 3703 (1) © 2013 Magnolia Press
The present list concerns the current classification and described diversity of the order Hymenoptera which
includes all wasps, bees and ants. Classification of extant superfamilies follows Sharkey (2007) and Sharkey
et al. (2011). The number of described and here included genus-group and species-group taxa are, unless
otherwise stated, updated until April 2012, while family-group taxa are updated until August 2013. In the
counts for fossil genera and species, only extinct taxa are included, not fossil representatives of extant taxa.
The taxa of Hymenoptera are divided into two suborders (Symphyta and Apocrita), with 27 superfamilies
(9 superfamilies in Symphyta and 18 in Apocrita) and 132 families. It is noteworthy that 17 of these families
have been described since year 2000 and that 16 of these newly recognized families represent now extinct
families only known from fossils. New genera and species are frequently encountered and our most recent
estimate total 8423 extant genera with an additional 685 extinct genera. Considered one of the most species-
rich insect orders, behind only Coleoptera and Lepidoptera, there is a total of 153088 extant and described
species, in addition to 2429 extinct described species.
Diversity estimates based on described species for some of the parasitic wasp families can be misleading.
There is a rich literature on the family Cynipidae with much ongoing research, but the classification is
currently changing and many new species are being described especially from the northern Neotropical and
Oriental regions. At the same time several groups remain without modern revisions, and there is a wealth of
synonyms particularly due to separate names of morphologically different generations in the subfamily
Cynipinae. Diversity counts for Figitidae and several other large families of parasitic Hymenoptera also bear
little relationship to the actual diversity. All major groups of Figitidae (most of the subfamily Eucoilinae, the
major genera in Charipinae, plus some genera of Figitinae, Anacharitinae and Aspicerinae) remain without
modern revisions, and there are very few current workers in the group. To exemplify this, of the 919 currently
formally valid species names in Eucoilinae, current researchers have examined less than 50% of the types and
assessed the generic combination—at most 400 species are confirmedly valid and actually recognizable. Even
in Europe, the historically best known area, at least 50% of the species present in collections are in fact still
undescribed, and for the rest of the world, this ratio is much higher (Forshage, pers. obs.). For real numbers of
species of Figitidae, the only figure someone has dared to estimate is an approximately 14,000 (Nordlander,
Order Hymenoptera Linnaeus, 1758 (2 suborders) (8,423 (and †684) genera, 152,677 (and †2,428) species)
Suborder Symphyta
(9 superfamilies, 25 families)
Superfamily Anaxyeloidea Martynov, 1925 (1 family)
Family Anaxyelidae Martynov, 1925 (1 (and †12) genera, 1 (and †32) species)
Superfamily Cephoidea Newman, 1834 (2 families)
Family Cephidae Newman, 1834 (21 (and †3) genera, 160 (and †6) species)
† Family Sepulcidae Rasnitsyn, 1968 (†15 genera, †38 species)
Superfamily Karatavitoidea Rasnitsyn, 1963 (1 family)
† Family Karatavitidae Rasnitsyn, 1963 (†5 genera, †6 species)
Superfamily Orussoidea Newman, 1834 (2 families)
Family Orussidae Newman, 1834 (16 (and †2) genera, 82 (and †3) species)
† Family Paroryssidae Martynov, 1925 (†4 genera, †10 species)
Superfamily Pamphilioidea Cameron, 1890 (3 families)
Family Megalodontesidae Konow, 1897 (1 (and †1) genera, 42 (and †1) species)
1. Taeger et al. 2010; Taeger & Blank 2012.
Zootaxa 3703 (1) © 2013 Magnolia Press · 53
Family Pamphiliidae Cameron, 1890 (10 (and †3) genera, 291 (and †5) species)
† Family Xyelydidae Rasnitsyn, 1986 (†8 genera, †20 species)
Superfamily Siricoidea Billberg, 1820 (6 families)
† Family Daohugoidae Rasnitsyn & Zhang, 2004 (†1 genus, †1 species)
† Family Praesiricidae Rasnitsyn, 1968 (†6 genera, †8 species)
† Family Protosiricidae Rasnitsyn & Zhang, 2004 (†1 genus, †1 species)
† Family Pseudosiricidae Handlirsch, 1908 (†1 genus, †14 species)
† Family Sinosiricidae Hong, 1975 (†1 genus, †1 species)
Family Siricidae Billberg, 1820 (11 (and †9) genera, 111 (and †13) species)
Superfamily Tenthredinoidea Latreille, 1803 (8 families)
Family Argidae Konow, 1890 (58 (and †1) genera, 897 (and †7) species)
Family Blasticotomidae Thomson, 1871 (2 (and †1) genera, 12 (and †1) species)
Family Cimbicidae W. Kirby, 1837 (16 (and †6) genera, 182 (and †19) species)
Family Diprionidae Rohwer, 1910 (11 (and †2) genera, 136 (and †2) species)
† Family Electrotomidae Rasnitsyn, 1977 (†1 genus, †1 species)
Family Pergidae Rohwer, 1911 (60 genera, 442 species)
Family Tenthredinidae Latreille, 1803 (400 (and †14) genera, 5500 (and †79) species)
† Family Xyelotomidae Rasnitsyn, 1968 (†14 genera, †21 species)
Superfamily Xiphydrioidea Leach, 1819 (1 family)
Family Xiphydriidae Leach, 1819 (28 genera, 146 species)
Superfamily Xyeloidea Newman, 1834 (1 families)
Family Xyelidae Newman, 1834 (5 (and †47) genera, 63 (and †93) species)
Suborder Apocrita (18 superfamilies, 105 families)
Superfamily Ephialtitoidea
Handlirsch, 1906 (1 family)
† Family Ephialtitidae Handlirsch, 1906 (†2 genera, †60 species)
Superfamily Trigonaloidea
Cresson, 1887 (2 families)
† Family Maimetshidae
Rasnitsyn, 1975 (†9 genera, †11 species)
Family Trigonalidae
Cresson, 1887 (16 (and †4) genera, 92 (and †5) species)
Superfamily Megalyroidea Schletterer, 1889 (1 family)
Family Megalyridae
Schletterer, 1889 (8 (and †11) genera, 43 (and †40) species)
Superfamily Stephanoidea Leach, 1815 (1 family)
Family Stephanidae
Leach, 1815 (11 (and †4) genera, 342 (and †8) species)
Superfamily Ceraphronoidea Haliday, 1833 (4 families)
Family Ceraphronidae
Haliday, 1833 (15 genera, 304 (and †2) species)
Family Megaspilidae
Ashmead, 1893 (12 (and †1) genera, 299 (and †12) species)
3. This name has been spelled in two ways: Trigonalidae and Trigonalyidae. The family was proposed as Trigonalidae by Cresson
(1887) and is based on the genus Trigonalys Westwood (1835). Westwood did not provide the derivation of his genus-group
name, but it must be assumed that Trigonalys is to be treated as a noun in the nominative singular. It is presumably a compound
word. If Latin, the first part of the name “trigonium” is a noun meaning triangle but the second part is ambiguous; it might come
from the Greek “alys” or it might be an arbitrary combination of letters. Only in the latter case does ICZN Article 29.3.3 apply
and the stem adopted by Cresson, who establishes the new family-group taxon, must be accepted as the correct spelling, namely
4. Perrichot et al. 2011.
5. Lelej 2003a; Poinar 2005.
6. Rasnitsyn 1975; Belokobylskij 1998; Perrichot 2009.
7. Aguiar 2004a, 2004b, 2005, 2006; Aguiar & Jennings 2005, 2010; Aguiar et al. 2010; Achterberg & Yang 2004; Achterberg &
Quicke 2006; Engel & Grimaldi 2004; Engel 2005b; Hong et al. 2010, 2011; Hong & Xu 2011.
8. Johnson & Musetti 2004; Evans et al. 2005.
9. Dessart 2001; Martinez 2003; Takada 2009; Peñalver & Engel 2006.
54 · Zootaxa 3703 (1) © 2013 Magnolia Press
† Family Radiophronidae
Ortega-Blanco, Rasnitsyn & Declòs, 2010 (†2 genera, †2 species)
† Family Stigmaphronidae
Kozlov, 1975 (†8 genera, †25 species)
Superfamily Evanioidea Latreille, 1802 (5 families)
† Family Andreneliidae
Rasnitsyn & Martínez-Declòs, 2000 (†1 genus, †1 species)
Family Aulacidae
Shuckard, 1841 (2 (and †10) genera, 185 (and †18) species)
Family Evaniidae
Latreille, 1802 (21 (and †9) genera, 449 (and †19) species)
Family Gasteruptiidae
Ashmead, 1900 (6 genera, 496 species)
† Family Praeaulacidae
Rasnitsyn, 1972 (†10 genera, †39 species)
Superfamily Mymarommatoidea Debauche, 1948 (3 families)
† Family Alavarommatidae
Ortega-Blanco, Peñalver, Delclòs & Engel, 2011 (†1 genus, †1
† Family Gallorommatidae
Gibson, Read & Huber, 2007 (†1 genus, †3 species)
Family Mymarommatidae
Debauche, 1948 (3 (and †2) genera, 10 (and †10) species)
Superfamily Proctotrupoidea Latreille, 1802 (11 families)
Family Austroniidae
Kozlov, 1975 (1 (and †1) genera, 3 (and †1) species)
Family Heloridae Förster, 1856 (1 (and †9) genera, 12 (and †13) species)
† Family Iscopinidae Rasnitsyn, 1980 (†10 genera, †39 species)
† Family Jurapriidae Rasnitsyn, 1983 (†1 genus, †1 species)
† Family Mesoserphidae
Kozlov, 1970 (†16 genera, †33 species)
Family Pelecinidae
Haliday, 1839 (1 (and †5) genera, 3 (and †8) species)
Family Peradeniidae
Naumann & Masner, 1985 (1 genus, 2 (and †1) species)
Family Proctorenyxidae
Lelej & Kozlov, 1999 (2 genera, 2 species)
Family Proctotrupidae Latreille, 1802 (28 (and †15) genera, 403 (and †26) species)
Family Roproniidae Bradley, 1905 (2 (and †4) genera, 20 (and †5) species)
Family Vanhorniidae Crawford, 1909 (3 genera, 5 species)
† Genera unplaced to family (†2 genera, †3 species)
Superfamily Diaprioidea Haliday, 1833 (5 families)
Family Diapriidae
Haliday, 1833 (190 (and †7) genera, 2048 (and †22) species)
Family Ismaridae
Thomson, 1858 (1 genus, 29 species)
Family Maamingidae
Early, Masner, Naumann & Austin, 2001 (1 genus, 2 species)
Family Monomachidae
Ashmead, 1902 (2 genera, 30 species)
10. Ortega-Blanco et al. 2010.
11. McKellar & Engel 2011.
12. Rasnitsyn & Martínez-Declòs 2000.
13. Turrisi et al. 2009; Jennings & Krogmann 2009.
14. Deans 2005; Kawada & Azevedo 2007; Deans & Kawada 2008; Rasnitsyn & Brothers 2007; Zhang & Rasnitsyn 2008; Peñalver
et al. 2010; Deans et al. 2012.
15. Jennings & Austin 2002; Macedo 2009.
16. Zhang & Rasnitsyn 2008.
17. Ortega-Blanco et al. 2011a.
18. Engel & Grimaldi 2007a.
19. Gibson et al. 2007; Huber et al. 2008.
20. Kozlov in Rasnitsyn 1975.
22. Engel & Grimaldi 2006a.
23. Johnson et al. 2001.
24. Lelej & Kozlov 1999; He et al. 2002.
25. Perrichot & Nel 2008.
27. Early et al. 2001.
28. Johnson & Musetti 2012.
Zootaxa 3703 (1) © 2013 Magnolia Press · 55
† Family Spathiopterygidae
Engel & Ortega-Blanco, 2013 (†3 genera and †3 species)
Superfamily Platygastroidea Haliday, 1833 (1 family)
Family Platygastridae Haliday, 1833 (236 (and †20) genera, 5385 (and †45) species)
Superfamily Serphitoidea Brues, 1937 (1 family)
† Family Serphitidae
Brues, 1937 (†3 genera, †9 species)
Superfamily Cynipoidea Latreille, 1802 (8 families)
Family Austrocynipidae
Riek, 1971 (1 genus, 1 species)
Family Cynipidae
Latreille, 1802 (74 (and †3) genera, 1412 (and †11) species)
Family Figitidae Hartig, 1840 (148 (and †10) genera, 1571 (and †11) species)
† Family Gerocynipidae
Liu & Engel, 2007 (†3 genera, †5 species)
Family Ibaliidae
Thomson, 1862 (3 (and †1) genera, 20 (and †2) species)
Family Liopteridae Ashmead, 1895 (10 (and †2) genera, 153 (and †2) species)
† Family Protimaspidae
Liu & Engel, 2007 (†1 genus, †1 species)
† Family Stolamissidae
Liu & Engel, 2007 (†1 genus, †1 species)
Superfamily Chalcidoidea
Latreille, 1817 (23 families)
Family Agaonidae
Walker, 1846 (20 (and †2) genera, 762 (and †4) species)
Family Aphelinidae Thomson, 1876 (29 (and †1) genera, 1078 (and †1) species)
Family Azotidae
Nikol’skaya & Yasnosh, 1966 (1 genus, 92 species)
Family Chalcididae Latreille, 1817 (90 (and †3) genera, 1469 (and †5) species)
Family Cynipencyrtidae
Trjapitzin, 1973 (1 genus, 1 species)
Family Encyrtidae Walker, 1837 (493 (and †2) genera, 4058 (and †3) species)
Family Eriaporidae
Ghesquière, 1955 (5 genera, 22 species)
Family Eucharitidae Walker, 1846 (57 (and †1) genera, 427 (and †1) species)
Family Eulophidae Westwood, 1829 (334 (and †2) genera, 4969 (and †3) species)
Family Eupelmidae Walker, 1833 (51 (and †4) genera, 931 (and †5) species)
Family Eurytomidae Walker, 1832 (97 (and †1) genera, 1453 (and †3) species)
† Family Khutelchalcididae Rasnitsyn, Basibuyuk & Quicke, 2004 (†1 genus, †1 species)
Family Leucospidae
Walker, 1834 (4 (and †1) genera, 134 (and †1) species)
Family Mymaridae Haliday, 1833 (96 (and †13) genera, 1437 (and †20) species)
Family Ormyridae Förster, 1856 (3 genera, 125 species)
Family Perilampidae Förster, 1856 (17 (and †1) genera, 284 (and †1) species)
Family Pteromalidae Dalman, 1820 (619 (and †6) genera, 3544 (and †20) species)
Family Rotoitidae
Bouček & Noyes, 1987 (2 genera, 2 species)
Family Signiphoridae Howard, 1894 (4 genera, 78 species)
Family Tanaostigmatidae Ashmead, 1904 (9 (and †1) genera, 93 (and †1) species)
29. Engel et al. 2013.
30. Ortega-Blanco et al. 2011b.
31. Ronquist 1999.
32. Liljeblad 2002.
33. Liu et al. 2007a.
34. Liu & Nordlander 1994; Liu et al. 2007; Liu & Engel 2010.
35. Liu et al. 2007a.
36. Liu et al. 2007a.
37. Noyes 2012.
38. Peñalver et al. 2006.
39. Heraty et al. (2013) revised the status of this family. The included taxa were previously found in Aphelinidae.
40. Heraty et al. (2013) revised the status of this family. The included taxa were previously found in Tanaostigmatidae.
41. Heraty et al. (2013) revised the status of this family. The included taxa were previously found in Aphelinidae.
42. Engel 2002.
43. Gibson & Huber 2000.
56 · Zootaxa 3703 (1) © 2013 Magnolia Press
Family Tetracampidae Förster, 1856 (15 (and †5 (+†1 from Mymaridae, unpubl.) genera, 44
(and †8 (+†1 from Mymaridae, unpublished) species)
Family Torymidae Walker, 1833 (82 (and †5) genera, 900 (and †13) species)
Family Trichogrammatidae Haliday, 1851 (97 (and †3) genera, 881 (and †3) species)
Superfamily Ichneumonoidea Latreille, 1802 (3 families)
Family Braconidae Nees, 1811 (1057 (and †5) genera, 19205 (and †206) species)
Family Ichneumonidae
Latreille, 1802 (1575 genera, 24025 (and †216) species)
† Family Praeichneumonidae
Rasnitsyn, 1983 (†1 genus, †5 species)
Superfamily Bethylonymoidea Rasnitsyn, 1975 (1 family)
† Family Bethylonymidae
Rasnitsyn, 1975 (†2 genera, †17 species)
Superfamily Chrysidoidea Latreille, 1802 (9 families)
Family Bethylidae
Haliday, 1839 (84 (and †11) genera, 2340 (and †13) species)
Family Chrysididae
Latreille, 1802 (81 (and †6) genera, 2500 (and †9) species)
Family Dryinidae
Haliday, 1833 (41 (and †2) genera, 1605 (and †19) species)
Family Embolemidae
Förster, 1856 (2 (and †1) genera, 39 (and †9) species)
† Family Falsiformicidae
Rasnitsyn, 1975 (†1 genus, †1 species)
† Family Plumalexiidae
Brothers, 2011 (†1 genus, †1 species)
Family Plumariidae
Bischoff, 1914 (7 genera, 22 species)
Family Sclerogibbidae
Ashmead, 1902 (3 (and †1) genera, 20 (and †2) species)
Family Scolebythidae
Evans, 1963 (4 (and †5) genera, 6 (and †7) species)
Superfamily Vespoidea
Latreille, 1802 (10 families)
Family Bradynobaenidae
de Saussure, 1892 (10 genera, 188 species)
Family Formicidae
Latreille, 1802 (299 (and †132) genera, 12199 (and †620) species)
Family Mutillidae
Latreille, 1802 (210 (and †1) genera, 4302 (and †12) species)
Family Pompilidae
Latreille, 1804 (125 (and †2) genera, 4855 (and †16) species)
Family Rhopalosomatidae
Ashmead, 1896 (4 (and †4) genera, 72 (and †4) species)
Family Sapygidae
Latreille, 1810 (12 (and †1) genera, 66 (and †1) species)
Family Scoliidae
Latreille, 1802 (143 (and †5) genera, 560 (and †17) species)
Family Sierolomorphidae
Krombein, 1951 (2 genera, 11 (and †1) species)
Family Tiphiidae
Leach, 1815 (120 (and †5) genera, 2000 (and †17) species)
Family Vespidae
Latreille, 1802 (268 (and †3) genera, 4932 (and †11) species)
44. Wahl 1999; Yu et al. 2005.
45. Kopylov 2012.
46. Rasnitsyn 1975; Rasnitsyn & Zhang 2010.
47. Vargas-Rojas & Terayama 2006; Huber 2009; Carpenter 1992; Ploëg & Nel 2004.
48. Kimsey & Bohart 1990; Engel 2006.
49. Olmi & Virla 2006; Engel 2005a; Olmi et al. 2011a.
50. Olmi 2006a; Olmi et al. 2011b; Perrichot & Engel 2011; Ortega-Blanco et al. 2011c.
51. Rasnitsyn 1975.
52. Brothers 2011.
53. Huber 2009; Brothers 2011.
54. Olmi 2006b; Engel & Grimaldi 2006b.
55. Brothers 2006; Engel & Grimaldi 2007b.
56. This equals Scolioidea Latreille, 1802 + Pompiloidea Latreille, 1804 + Vespoidea Latreille, 1802 + Formicoidea Latreille, 1802
sensu Rasnitsyn 1988.
57. Lelej 2003b.
58. (accessed March 2012).
59. Brothers 2003; Lelej 2005.
60. Engel & Grimaldi 2006c.
61. (accessed August 2013).
62. Kurzenko 2009; Bennett & Engel 2005.
Zootaxa 3703 (1) © 2013 Magnolia Press · 57
Superfamily Apoidea Latreille, 1802 (13 families)
Family Ampulicidae
Shuckard, 1840 (6 (and †7) genera, 200 (and †8) species)
Family Andrenidae
Latreille, 1802 (77 (and †3) genera, 2917 (and †11) species)
† Family Angarosphecidae
Rasnitsyn, 1975 (†13 genera, †44 species)
Family Apidae
Latreille, 1802 (209 (and †22) genera, 5749 (and †87) species)
Family Colletidae
Lepeletier de Saint Fargeau, 1841 (86 genera, 2547 (and †2) species)
Family Crabronidae
Latreille, 1802 (242 (and †24) genera, 8773
(and †29) species)
Family Halictidae
Thomson, 1869 (79 (and †7) genera, 4327 (and †22) species)
Family Megachilidae
Latreille, 1802 (76 (and †6) genera, 4096 (and †34) species)
Family Melittidae
Schenck, 1860 (15 (and †2) genera, 187 (and †4) species)
† Family Mellitosphecidae
Poinar & Danforth, 2006 (†1 genus, †1 species)
† Family Paleomelittidae
Engel, 2001 (†1 genus, †1 species)
Family Sphecidae
Latreille, 1802 (19 (and †1) genera, 724
(and †1) species)
Family Stenotritidae
Cockerell, 1934 (2 genera, 21 species)
Apocrita incertae sedis (3 families)
† Family Archaeocynipidae
Rasnitsyn & Kovalev, 1988 (†2 genera, †3 species)
† Family Eostephanitidae Hong, 2002 (†1 genus, †1 species)
† Family Kuafuidae
Rasnitsyn & Zhang, 2010 (†3 genera, †3 species)
Gerardo Lamas and Josef Straka are thanked for information about the spelling of Trigonalidae. Support to C.
Rasmussen was provided by a Carlsberg Foundation grant, support to A.P. Aguiar was provided by research
funding from FAPES – Fundação de Amparo à Pesquisa do Espírito Santo (Process 45.440.611/2009), support
to A.S. Lelej was provided by Far Eastern Branch of the Russian Academy of Sciences (# 12-I-П-30-03),
support to N.F. Johnson was supported in part by the National Science Foundation under grant No. DEB-
0614764 to N.F. Johnson and A.D. Austin and the Evanioidea contribution was funded, in part, by the U.S.
National Science Foundation (award DEB-0842289).
63. Osten (2005) ignored Argaman's classification and lists only 43 valid genera for Scoliidae. While Argaman's division of the
family into 143 genera is not supported by phylogenetic analysis or proper diagnosis, the family obviously is very diverse and in
need of a generic revision based on a phylogeny.
64. Argaman 1996; Osten 2005; Zhang 2006.
65. Gorbatovskij & Lelej 1998; Rasnitsyn 2000.
66. Kimsey 1991; Kimsey 2006; Engel et al. 2009.
67. Carpenter & Rasnitsyn 1990; Sarmiento & Carpenter 2006; Brothers & Rasnitsyn 2008.
69. Michener 2007; Michez et al. 2012; Ascher & Pickering 2012.
71. Michener 2007; Michez et al. 2012; Ascher & Pickering 2012.
72. Michener 2007; Michez et al. 2012; Ascher & Pickering 2012.
74. Updated August 2013.
75. Michener 2007; Michez et al. 2012; Ascher & Pickering 2012.
76. Michener 2007; Michez et al. 2012; Ascher & Pickering 2012.
77. Michener 2007; Michez et al. 2012; Ascher & Pickering 2012.
78. Michez et al. 2012.
79. Michener 2007; Michez et al. 2012; Ascher & Pickering 2012.
81. Updated August 2013.
82. Michener 2007; Ascher & Pickering 2012.
58 · Zootaxa 3703 (1) © 2013 Magnolia Press
Achterberg, C. van & Quicke, D.L.J. (2006) Taxonomic notes on Old World Stephanidae (Hymenoptera): description of
Parastephanellus matsumotoi sp. n. from Japan, redescription of Commatopus xanthocephalus (Cameron) and keys
to the genera Profoenatopus van Achterberg and Megischus Brullé. Tijdschrift voor Entomologie, 149, 215–225.
Achterberg, C. van & Yang, Z.Q. (2004) New species of the genera Megischus Brullé and Stephanus Jurine from China
(Hymenoptera: Stephanoidea: Stephanidae), with a key to world species of the genus Stephanus. Zoologische
Medelingen Leiden, 78, 101–117.
Aguiar, A.P. (2004a) World catalog of the Stephanidae (Hymenoptera: Stephanoidea). Zootaxa, 753, 1–120.
Aguiar, A.P. (2004b) Additions to the revision of the genus Hemistephanus Enderlein (Hymenoptera: Stephanidae), with
inclusion of four taxa and description of two new species. Papéis Avulsos de Zoologia, 44, 13–43.
Aguiar, A.P. (2005) A New and Unusual Species of Stephanidae (Hymenoptera), with a Discussion on its Phylogenetic
Implications. Journal of Hymenoptera Research, 14, 1–6.
Aguiar, A.P. (2006) The Stephanidae (Hymenoptera) of Mexico, with description of six new species and key to western
Foenatopus Smith. Zootaxa, 1186, 1–56.
Aguiar, A.P. & Jennings, J.T. (2005) First record of Stephanidae (Hymenoptera) from New Caledonia, with descriptions
of four new species of Parastephanellus Enderlein. Zootaxa, 1001, 1–16.
Aguiar, A.P. & Jennings, J.T. (2010) Order Hymenoptera, family Stephanidae. In: van Harten, A. (Ed.), Arthropod fauna
of the United Arab Emirates. Vol. 3. Dar Al Ummah Printing, Publishing, Distribution & Advertising, Abu Dhabi,
pp. 299–305.
Aguiar, A.P., Jennings, J.T. & Turrisi, G.F. (2010) Three new Middle–Eastern species of Foenatopus Smith
(Hymenoptera: Stephanidae) with a new host record and key to species with two spots on the metasoma. Zootaxa,
2714, 40–58.
Argaman Q. (1996) Generic synopsis of Scoliidae (Hymenoptera, Scoloidea). Annales Historico–Naturales Musei
Nationalis Hungarici, 88, 171–222.
Ascher, J.S. & Pickering, J. (2012) Discover Life bee species guide and world checklist (Hymenoptera: Apoidea:
Anthophila). (accessed February 20, 2012).
Belokobylskij, S.A. (1998) Fam. Megalyridae. In: Lehr P.A. (Ed.), [Key to the Insects of Russian Far East, Vol. 4, pt. 3].
Dalnauka, Vladivostok, pp. 657–659. (In Russian).
Bennett, D. J. & Engel, M.S. (2005) A primitive sapygid wasp in Burmese amber (Hymenoptera: Sapygidae). Acta
zoologica cracoviensia, 48B, 1–9.
Brothers, D.J. (2003) The first fossil Ephutini (Hymenoptera: Mutillidae), a new species of Ephuta Say from Dominican
amber. Acta zoologica cracoviensia, 46 (suppl. - Fossil Insects), 101–107.
Brothers, D.J. (2006) Familia Scolebythidae. In: Fernández, F. & Sharkey, M.J. (Eds.), Introducción a los Hymenoptera
de la Región Neotropical. Sociedad Colombiana de Entomología y Universidad Nacional de Colombia, Bogotá,
D.C., pp. 391–392.
Brothers, D.J. (2011) A new Late Cretaceous family of Hymenoptera, and phylogeny of the Plumariidae and
Chrysidoidea (Aculeata). ZooKeys, 130, 515–542.
Brothers, D.J. & Rasnitsyn, A.P. (2008) A new genus and species of Euparagiinae from the Late Cretaceous of southern
Africa (Hymenoptera: Vespidae). Alavesia, 2, 73–76.
Carpenter F.M. (1992) Superclass Hexapoda. In: Kaesler, R.I. (Ed.), Treatise on Invertebrate Paleontology, Part R
Arthropoda 3–4. Geological Society America, Boulder, Colorado, pp. 1–655.
Carpenter, J.M. & Rasnitsyn, A.P. (1990) Mesozoic Vespidae. Psyche, 97, 1–20.
Cresson, E.T. (1887) Synopsis of the families and genera of the Hymenoptera of America, North of Mexico, together
with a catalogue of the described species, and bibliography. Transactions of the American Entomological Society,
Supplementary volume, i–vi, 1–350.
Deans, A.R. (2005) Annotated catalog of the world's ensign wasp species (Hymenoptera: Evaniidae). Contributions of
the American Entomological Institute, 34, 1–164.
83. Rasnitsyn & Kovalev 1988.
84. Rasnitsyn & Zhang 2010.
Zootaxa 3703 (1) © 2013 Magnolia Press · 59
Deans, A.R. & Kawada, R. (2008) Alobevania, a new genus of Neotropical ensign wasps (Hymenoptera: Evaniidae),
with three new species: integrating taxonomy with the World Wide Web. Zootaxa, 1787, 28–44.
Deans, A.R., Yoder, M.J. & Dole, K. (2012) Evanioidea Online - catalog of information about evanioid wasps
(Hymenoptera). (accessed 20.03.2012).
Dessart, P. (2001) Les Megaspilinae ni européens, ni américains 2. Les Dendrocerus Ratzeburg, 1852, à mâles non
flabellicornés (Hymenoptera Ceraphronoidea Megaspilidae). Belgian Journal of Entomology, 3, 3–124.
Early, J.W., Masner, L., Naumann, I.D. & Austin, A.D. (2001) Maamingidae, a new family of proctotrupoid wasp
(Insecta: Hymenoptera) from New Zealand. Invertebrate Taxonomy, 15, 341–352.
Engel, M.S. (2002) The first leucospid wasp from the fossil record (Hymenoptera: Leucospidae). Journal of Natural
History, 36, 435–441.
Engel, M.S. (2005a) A dryinine wasp in Burmese amber (Hymenoptera: Dryinidae). Polskie Pismo Entomologiczne, 74,
Engel, M.S. (2005b) The crown wasp genus Electrostephanus (Hymenoptera: Stephanidae): Discovery of the female and
a new species. Polskie Pismo Entomologiczne, 74, 317–332.
Engel, M.S. (2006) A new cuckoo wasp of the genus Ceratochrysis in amber from the Dominican Republic
(Hymenoptera: Chrysididae). Polskie Pismo Entomologiczne, 75, 499–504.
Engel, M.S. & Grimaldi, D.A. (2004) The first Mesozoic stephanid wasp (Hymenoptera: Stephanidae). Journal of
Paleontology, 78, 1192–1197.
Engel, M.S. & Grimaldi, D.A. (2006a) A diminutive pelecinid wasp in Cretaceous amber from New Jersey
(Hymenoptera: Pelecinidae). Northeastern Naturalist, 13, 291–297.
Engel, M.S. & Grimaldi, D.A. (2006b) The first Cretaceous sclerogibbid wasp (Hymenoptera: Sclerogibbidae).
American Museum Novitates, 3515, 1–7.
Engel, M.S. & Grimaldi, D.A. (2006c) The first Cretaceous spider wasp (Hymenoptera: Pompilidae). Journal of the
Kansas Entomological Society, 79, 359–368.
Engel, M.S. & Grimaldi, D.A. (2007a) New false fairy wasps in Cretaceous amber from New Jersey and Myanmar
(Hymenoptera: Mymarommatoidea). Transactions of the Kansas Academy of Science, 110, 159–168.
Engel, M.S. & Grimaldi, D.A. (2007b) Cretaceous Scolebythidae and phylogeny of the family (Hymenoptera:
Chrysidoidea). American Museum Novitates, 3568, 1–16.
Engel, M.S., Ortega-Blanco J. & Bennett D.J. (2009) A remarkable tiphiiform wasp in mid–Cretaceous amber from
Myanmar (Hymenoptera: Tiphiidae). Transactions of the Kansas Academy of Science, 112, 1–6.
Engel, M.S., Ortega-Blanco, J., Soriano, C., Grimaldi, D.A., & Martinez-Delclos, X. (2013) A new lineage of enigmatic
diaprioid wasps in Cretaceous amber (Hymenoptera, Diaprioidea). American Museum Novitates, 3771, 1–23.
Evans, G.A., Dessart, P. & Glenn, H. (2005) Two new species of Aphanogmus (Hymenoptera: Ceraphronidae) of
economic importance reared from Cybocephalus nipponicus (Coleoptera: Cybocephalidae). Zootaxa, 1018, 47–54.
Gibson, G.A.P. & Huber, J.T. (2000) Review of the family Rotoitidae (Hymenoptera; Chalcidoidea), with description of
new genus and species from Chile. Journal of Natural History, 34, 2293–2314.
Gibson, G.A.P., Read, J. & Huber, J.T. (2007) Diversity, classification and higher relationships of Mymarommatoidea
(Hymenoptera). Journal of Hymenoptera Research, 16, 51–146.
Gorbatovskij, V.V. & Lelej, A.S. (1998) Fam. Sierolomorphidae. In: Lehr P.A. (Ed.), [Key to the Insects of Russian Far
East, Vol. 4, pt. 3]. Dalnauka, Vladivostok, pp. 683–684. (In Russian).
He, J.-H., Ma, Y. & Chen, X.-X. (2002) A new record of Proctorenyxidae from China (Hymenoptera: Proctotrupoidea).
Acta Zootaxonomia Sinica, 27, 630. (In Chinese).
Heraty, J.M., Burks, R.A., Cruaud, A., Gibson, G.A.P., Liljeblad, J., Munro, J., Rasplus, J.-Y., Delvare, G., Janšta, P.,
Gumovsky, A., Huber, J.T., Woolley, J.B., Krogmann, L., Heydon, S.L., Polaszek, A., Schmidt, S., Darling, D.C.,
Gates, M.W., Mottern, J., Murray, E., Molin, A.D., Triapitsyn, S., Baur, H., Pinto, J.D., Noort, S.v., George, J., &
Yoder, M.J. (2013) A phylogenetic analysis of the megadiverse Chalcidoidea (Hymenoptera). Cladistics.
Hong, C.D., Achterberg, C. van & Xu, Z.F. (2010) A new species of Megischus Brullé (Hymenoptera, Stephanidae) from
China, with a key to the Chinese species. ZooKeys, 69, 59–64.
Hong, C.D., Achterberg, C. van & Xu, Z.F. (2011) A revision of the Chinese Stephanidae (Hymenoptera, Stephanoidea).
ZooKeys, 110, 1–108.
Hong, C.D. & Xu, Z.F. (2011) A newly recorded genus and species of family Stephanidae (Hymenoptera, Stephanoidea)
from China. Entomotaxonomia, 33, 71–73.
60 · Zootaxa 3703 (1) © 2013 Magnolia Press
Huber, J.T. (2009) Biodiversity of Hymenoptera. In: Foottit, R.G. & Adler, P.H. (Eds.), Insect biodiversity: science and
society. Blackwell Publishing, pp. 303–323.
Huber, J.T., Gibson, G.A.P., Bauer, L.S., Liu, H. & Gates, M. (2008) The genus Mymaromella (Hymenoptera:
Mymarommatidae) in North America, with a key to described extant species. Journal of Hymenoptera Research, 17,
International Commission on Zoological Nomenclature (ICZN). (1999) International Code of Zoological Nomenclature.
Fourth Edition. ICZN, London . i–xxx + 306 pp.
Jennings, J.T. & Austin, A.D. (2002) Systematics and distribution of world hyptiogastrine wasps (Hymenoptera:
Gasteruptiidae). Invertebrate Systematics, 16, 735–811.
Jennings, J.T. & Krogmann, L. (2009) A new species of Pristaulacus Kieffer (Hymenoptera; Aulacidae) from Baltic
amber. Insect Systematics & Evolution, 40, 201–207.
Johnson, N.F. & Musetti, L. (2004) Catalog of the systematic literature of the superfamily Ceraphronoidea
Contributions of the American Entomological Institute, 33, 1–149.
Johnson, N.F. & Musetti, L. (2012) Genera of the parasitoid wasp family Monomachidae (Hymenoptera: Diaprioidea).
Zootaxa, 3188, 31–41.
Johnson, N.F., Musetti, L. & Janzen, J.-W. (2001) A new fossil species of the Australian endemic genus Peradenia
Naumann & Masner (Hymenoptera: Proctotrupoidea, Peradeniidae) from Baltic Amber. Insect Systematics &
Evolution, 32, 191–194.
Kawada, R. & Azevedo, C.O. (2007) Taxonomic revision of the Neotropical ensign wasp genus Decevania
(Hymenoptera: Evaniidae). Zootaxa, 1496, 1–30.
Kimsey, L.S. (1991) Relationships among the tiphiid wasp subfamilies (Hymenoptera). Systematic Entomology, 16, 427–
Kimsey, L.S. (2006) Familia Tiphiidae. In: Hanson, P.E. & Gauld I.D. (Eds.), Hymenoptera de la Región Neotropical.
The American Entomological Institute, Gainesville, pp. 575–583.
Kimsey, L.S. & Bohart, R. (1990) The Chrysidid Wasps of the World. Oxford University Press, Oxford, 652 pp.
Kopylov, D.S. (2012) New species of Praeichneumonidae (Hymenoptera, Ichneumonoidea) from the Lower Cretaceous
of Transbaikalia. Paleontological Journal, 46, 66–72.
Kurzenko, N.V. (2009) [Fam. Sapygidae–sapygids]. In: Storozhenko S.Y. (Ed.), [Insects of Lazovsky nature reserve].
Dalnauka, Vladivostok, pp. 224–225. (In Russian).
Lelej, A.S. (2003a) A review of the family Trigonalyidae (Hymenoptera) of the Palaearctic region. Far Eastern
Entomologist, 130, 1–7.
Lelej, A.S. (2003b) Review: G. Pagliano. Revisione della sottofamilia Apterogyninae (Hymenoptera: Bradynobaenidae)
Museo Regionale di Scienze Naturali. Torino, 2002, 387 pp. Entomologicheskoe obozrenie, 82, 526–527. (In
Lelej, A.S. (2005) Catalogue of the Mutillidae (Hymenoptera) of the Oriental Region. Dalnauka, Vladivostok, 1–252.
Lelej, A.S. & Kozlov, M. A. (1999) Proctorenyxidae nom. n. and Proctorenyxa nom. n. a new replacement names for
Renyxidae Kozlov and Renyxa Kozlov (Hymenoptera, Proctotrupoidea). Far Eastern Entomologist, 74, 6–7.
Liljeblad, J. (2002) Phylogeny and evolution of gall wasps (Hymenoptera: Cynipidae). Dissertation Uppsala University,
Liu, Z. & Engel, M.S. (2010) Baltic amber Ibaliidae (Hymenoptera: Cynipoidea): a new genus with implications for the
phylogeny and historical biogeography of the family. Systematic Entomology, 35, 164–171.
Liu, Z., Engel, M.S. & Grimaldi, D. (2007a) Phylogeny and geological history of the cynipoid wasps (Hymenoptera:
Cynipoidea). American Museum Novitates, 3583, 1–48.
Liu, Z. & Nordlander, G. (1994) Review of the family Ibaliidae (Hymenoptera: Cynipoidea) with keys to genera and
species of the world. Entomologica Scandinavica, 25, 377–392.
Liu, Z., Ronquist, F. & Nordlander, G. (2007b) The cynipoid genus Paramblynotus, phylogeny and historical
biogeography. Bulletin of the American Museum of Natural History, 304, 1–151.
Macedo, A.C.C. (2009) Generic classification for the Gasteruptiinae (Hymenoptera: Gasteruptiidae) based on a cladistic
analysis, with the description of two new Neotropical genera and the revalidation of Plutofoenus Kieffer. Zootaxa,
2075, 1–32.
Zootaxa 3703 (1) © 2013 Magnolia Press · 61
Martinez, J.J. (2003) Una nueva especie de Dendrocerus (Hymenoptera: Megaspilidae) de La Pampa, Argentina. Revista
de la Sociedad Entomológica Argentina, 62, 65–68.
McKellar, R.C. & Engel, M.S. (2011) New Stigmaphronidae and Megaspilidae (Hymenoptera: Ceraphronoidea) from
Canadian Cretaceous amber. Cretaceous Research, 32, 794–805.
Michener, C.D. (2007) The Bees of the World, Second Edition. Johns Hopkins University Press, Baltimore, xvi+953.
Michez, D., Vanderplanck, M. & Engel, M.S. (2012) Fossil bees and their plant associates. In: Patiny, S. (Ed.), Evolution
of Plant–Pollinator Relationships. Cambridge University Press, Cambridge, pp. 103–164.
Nordlander, G. (1984) Vad vet vi om parasitiska Cynipoidea? Entomologisk Tidskrift, 105, 36–40.
Noyes, J. (2012) Universal Chalcidoidea Database. World Wide Web electronic publication.
chalcidoids (accessed March 2012).
Olmi, M. (2006a) Familia Embolemidae. In: Fernández, F. & Sharkey, M.J. (Eds.), Introducción a los Hymenoptera de la
Región Neotropical. Sociedad Colombiana de Entomología y Universidad Nacional de Colombia, Bogotá, D.C., pp.
Olmi, M. (2006b) Familia Sclerogibbidae. In: Fernández, F. & Sharkey, M.J. (Eds.), Introducción a los Hymenoptera de
la Región Neotropical. Sociedad Colombiana de Entomología y Universidad Nacional de Colombia, Bogotá, D.C.,
pp. 393–395.
Olmi, M., Guglielmino, A. & Vollaro, M. (2011a) Revision of fossil species of Dryinus belonging to constans group,
with description of a new species (Hymenoptera: Dryinidae). Zootaxa, 2981, 43–55.
Olmi, M., Rasnitsyn, A.P. & Guglielmino, A. (2011b) The first record of Embolemidae (Hymenoptera Chrysidoidea) in
the Rovno Amber (Upper Eocene) of Ukraine: a male of Ampulicomorpha succinalis Brues. Paleontological
Journal, 45, 73–76.
Olmi, M. & Virla, E. (2006) Familia Dryinidae. In: Fernández, F. & Sharkey, M.J. (Eds.), Introducción a los
Hymenoptera de la Región Neotropical. Sociedad Colombiana de Entomología y Universidad Nacional de
Colombia, Bogotá, D.C., pp. 401–418.
Ortega-Blanco, J., Delclòs, X. & Engel, M.S. (2011c) The wasp family Embolemidae in Early Cretaceous amber from
Spain (Hymenoptera: Chrysidoidea). Journal of the Kansas Entomological Society, 84, 36–42.
Ortega-Blanco, J., Delclòs, X., Peñalver, E. & Engel, M.S. (2011b) Serphitid wasps in Early Cretaceous amber from
Spain (Hymenoptera: Serphitidae). Cretaceous Research, 32, 143–154.
Ortega-Blanco, J,. Peñalver, E., Delclòs, X. & Engel, M.S. (2011a) False fairy wasps in Early Cretaceous amber from
Spain (Hymenoptera: Mymarommatoidea). Paleontology, 54, 511–523.
Ortega-Blanco, J., Rasnitsyn , A. P. & Delclòs, X. (2010) A new family of ceraphronoid wasps from Early Cretaceous
Álava Amber, Spain.
Acta Palaeontologica Polonica, 55, 265–276.
Osten, T. (2005) Checkliste der Dolchwespen der Welt (Hymenoptera: Scoliidae). 62. Bericht der Naturforschenden
Gesellschaft Augsburg, 220, 1–63.
Peñalver, E. & Engel, M.S. (2006) Two wasp families rare in the fossil record (Hymenoptera): Perilampidae and
Megaspilidae from the Miocene of Spain. American Museum Novitates, 3540, 1–12.
Peñalver, E., Engel, M.S. & Grimaldi, D.A. (2006) Fig wasps in Dominican amber (Hymenoptera: Agaonidae).
American Museum Novitates, 3541, 1–16.
Peñalver, E., Ortega-Blanco, J., Nel, A. & Delclòs, X. (2010) Mesozoic Evaniidae (Insecta: Hymenoptera) in Spanish
amber: reanalysis of the phylogeny of the Evanioidea. Acta Geologica Sinica, 84, 809–827.
Perrichot, V. (2009) Long-tailed wasps (Hymenoptera: Megalyridae) from Cretaceous and Paleogene European amber.
Paleontological Contributions, 1, 1–35.
Perrichot, V. & Engel, M.S. (2011) A new micropterous species of Embolemus Westwood from Baltic amber
(Hymenoptera: Embolemidae). Annales de Paleontologie, 97, 1–7.
Perrichot, V. & Nel, A. (2008) A new belytine wasp in Cretaceous amber from France (Hymenoptera: Diapriidae).
Alavesia, 2, 203–209.
Perrichot, V., Ortega-Blanco, J., McKellar, R.C., Delclòs, X., Azar, D., Nel, A., Tafforeau, P. & Engel, M.S. (2011) New
and revised maimetshid wasps from Cretaceous ambers (Hymenoptera, Maimetshidae). ZooKeys, 130, 421–453.
Ploëg, G. de & Nel, A. (2004) A new bethylid wasp from the Lowermost Eocene amber of France (Hymenoptera:
Bethylidae: Bethylinae). Geologica Acta, 2, 75–82.
Poinar, G. (2005) Fossil Trigonalidae and Vespidae (Hymenoptera) in Baltic amber. Proceedings of the Entomological
Society of Washington, 107, 55–63.
62 · Zootaxa 3703 (1) © 2013 Magnolia Press
Rasnitsyn, A.P. (1975) [Hymenoptera Apocrita of the Mesozoic]. [Transactions of the Paleontological Institute Academy
Sciences U. S. S. R. 147], 134 pp. (In Russian).
Rasnitsyn, A.P. (1988) An outline of evolution of the hymenopterous insects (order Vespida). Oriental Insects, 22, 115–
Rasnitsyn, A.P. (2000) An extremely primitive aculeate wasp in the Cretaceous amber from New Jersey (Vespida:
?Sierolomorphidae. In: Grimaldi, D.A. (Ed.), Studies on Fossils in Amber, with Particular Reference to the
Cretaceous of New Jersey. Backhuys Publishers, Leiden, pp. 327–332.
Rasnitsyn, A.P. & Brothers, D.J. (2007) Two new hymenopteran fossils from the mid-Cretaceous of southern Africa
(Hymenoptera: Jurapriidae, Evaniidae). African Invertebrates, 48, 193-202.
Rasnitsyn, A.P. & Kovalev, O.V. (1988) The oldest Cynipoidea (Hymenoptera: Archaeocynipidae fam. n.) from the Early
Cretaceous Transbaikalia. Vestnik Zoologii, 1,18–21. (In Russian).
Rasnitsyn, A.P. & Martínez-Declòs, X. (2000) Wasps (Insecta: Vespida = Hymenoptera) from the Early Cretaceous of
Spain. Acta Geologica Hispanica, 35, 65–95.
Rasnitsyn, A.P. & Zhang, H.-C. (2010) Early evolution of Apocrita (Insecta, Hymenoptera) as indicated by new findings
in the Middle Jurassic of Daohugou, Northeast China. Acta Geologica Sinica, 84, 834–873.
Ronquist, F. (1999) Phylogeny, classification and evolution of the Cynipoidea. Zoologica Scripta, 28, 139–164.
Sarmiento, C.E. & Carpenter, J.M. (2006) Familia Vespidae. In: Fernández, F. & Sharkey, M.J. (Eds.), Introducción a los
Hymenoptera de la Región Neotropical. Sociedad Colombiana de Entomología y Universidad Nacional de
Colombia, Bogotá, D.C., pp. 539–555.
Sharkey, M.J. (2007) Phylogeny and classification of Hymenoptera. Zootaxa, 1668, 521–548.
Sharkey, M.J., Carpenter, J.M., Vilhelmsen, L., Heraty, J., Liljeblad, J., Dowling, A.P.G., Schulmeister, S., Murray, D.,
Deans, A.R., Ronquist, F., Krogmann, L. & Wheeler, W.C. (2011) Phylogenetic relationships among superfamilies
of Hymenoptera. Cladistics, 27, 1–33.
Taeger, A. & Blank, S.M. (2012) ECatSym – Electronic World Catalog of Symphyta (Insecta, Hymenoptera) Program
version 3. 10, data version 38 (07.12.2011) Digital Entomological Information, Müncheberg. http://sdei. de/ecatsym/
(accessed 01.02.2012).
Taeger, A., Blank, S. & Liston, D. (2010) World Catalog of Symphyta (Hymenoptera). Zootaxa, 2580, 1–1064.
Takada, H. (2009), Description of a new Dendrocerus species (Hymenoptera: Megaspilidae) hyperparasitic on Stomaphis
aphids and additional notes on their primary parasitoid, Protaphidius nawaii (Braconidae), and another
hyperparasitoid, Euneura stomaphidis (Pteromalidae). Entomological Science, 12, 91–97.
Turrisi, G.F., Jennings, J.T. & Vilhelmsen, L. (2009) Phylogeny and generic concepts in the parasitoid wasp family
Aulacidae (Hymenoptera: Evanioidea). Invertebrate Systematics, 23, 27–59.
Vargas-Rojas, J.M. & Terayama, M. (2006) Familia Bethylidae. In: Fernández, F. & Sharkey, M.J. (Eds.), Introducción a
los Hymenoptera de la Región Neotropical. Sociedad Colombiana de Entomología y Universidad Nacional de
Colombia, Bogotá, D.C., pp. 427–442.
Wahl, D.B. (1999) Classification and Systematics of the Ichneumonidae (Hymenoptera). Available at http://hymfiles.
biosci. ohio–state. edu/catalogs/ichneumonids/ (accessed 15 February 2012).
Westwood, J.O. (1835) Specimens were exhibited of various Hymenopterous Insects, partly from the collection of the
Rev. F. W. Hope, and partly from that of Mr. Westwood. They were accompanied by characters by Mr. Westwood.
Proceedings of the Zoological Society of London, 3, 51–54.
Yu, D. S., Achterberg, C. & Horstmann, K. (2005) Interactive Catalogue of World Ichneumonoidea Taxonomy, biology,
morphology and distribution Compact Disc (Master version) Taxapad. Vancouver, Canada.
Zhang, H.-C., Rasnitsyn, A.P. (2008) Middle Jurassic Praeaulacidae (Insecta: Hymenoptera: Evanioidea) of Inner
Mongolia and Kazakhstan. Journal of Systematic Palaeontology, 6, 463–487.
Zhang, J.-F. (2006) A proscoliine wasp (Insecta: Hymenoptera: Scoliidae) from Shandong peninsula, East Asia.
Cretaceous Research, 27, 788–791.
... They are highly correlated with the diversity of plants and arthropods (Duelli & Obrist, 2003;Fabian et al., 2014;Guo et al., 2021), and therefore are useful biological indicators (Brock et al., 2021). They also include three guilds with different foraging types-pollinators, predators, and parasitoids-in a closely related taxonomic group (three superfamilies: Apoidea, Chrysidoidea, and Vespoidea; Aguiar et al., 2013). Their similarity facilitates the use of common traits and allows the observation of guild-specific responses without taxonomic features. ...
... The identification of the bee and wasp species was conducted as per the guidelines of Tadauchi and Murao (2014) and Terayama and Suda (2016), respectively. Family groups followed Aguiar et al. (2013). All the species were divided into three guilds: pollinators, predators, and parasitoids, according to the guidelines (Tadauchi & Murao, 2014;Terayama & Suda, 2016). ...
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The response of communities to environmental change is expected to vary among feeding guilds. To evaluate the response of guilds to environmental factors without considering the taxonomic specificities, it is useful to examine Aculeata bees and wasps, which consist of closely related taxa including different guilds, pollinators, predators, and parasitoids. In this study, we evaluated changes in species diversity (SD) and functional traits of each feeding guild along an elevational gradient in a boreal forest in northern Japan. We used yellow pan traps to collect Aculeata bees and wasps at 200–1600 m above sea level. We investigated six functional traits (trophic level, seasonal duration, body size, elevational range, nesting position, and soil dependency) and the horizontal distribution of the species. The SD of all Aculeata, predators, and parasitoids decreased with an increase in elevation; however, the SD of pollinators did not show any specific trend. Although the functional trait composition of all Aculeata species did not show any trend, that of each feeding guild responded to elevation in different ways. Pollinators increased in body size and showed a decrease in seasonal duration with increasing elevation, suggesting that tolerance and seasonal escape from physical stress at high elevations are important for shaping pollinator communities. Predators increased their elevational range and the proportion of above‐ground nesting species increased with increasing elevation, suggesting that the ability to live in a wider range of environments and avoid unsuitable soil environments at high elevations might be important. Parasitoids changed their hosts and displayed variable traits with increasing elevation, suggesting that brood parasitoids have difficulty in surviving at high elevation. The traits for each guild responded in different ways, even if they were dominated by the same environmental factors. Our findings imply that differences in the responses of functional traits would produce different community assembly patterns in different guilds during further climate change. The response of communities to climate change is expected to vary among feeding guilds. We found that functional trait composition of each feeding guild in Aculeata bees and wasps community responded to the elevational gradient in different ways. This differences can produce different community assembly patterns in different guilds during further climate change.
... Among the order Hymenoptera, the superfamily Chalcidoidea is extremely important in the control of various pests (Noyes, 2020). About 22,785 chalcidoid species have been identified belonging to 2,117 genera, constituting about one-third of all known parasitoid wasps (Aguiar et al., 2013;Noyes, 2020). More than 800 species in this group have been used in biological control programs (Noyes, 2020). ...
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Chalcidoidea play an important role in the biological control of many agricultural pests. The present study determines and compares the diversities of this superfamily in four ecosystems in the Lake Urmia basin. It is the largest hypersaline lake in the world with a large basin of different ecological conditions in northwest Iran. These stations were: Khajeh (K) and Tasuj (T) in Lake Urmia Basin (LUBin) and Khoda-Afarin (KA) and Tikme-Dash (TD) out of their (LUBout). Four months of Malaise trapping in these habitats resulted in a total of 512 specimens from 11 families of Chalcidoidea, including 6, 9, 10 and 10 families in K, KA, T and TD, respectively. Maximum abundance was observed in the T station with 192 and the minimum was in the K station with 38. However, the families, Encyrtidae and Chalcididae have maximum abundance in T and KA, respectively. The maximum activity of chalcidoid wasps was observed during June-August. The majority of the species were belonging to the family Mymaridae (25%), in LUB out habitats peaked at 47 specimens in TD during the sampling period, while the corresponding LUBin habitats peaked at 40 specimens in T. The two LUBout habitats had greater community similarity (60%) than found between in and LUB out habitats (42%) and diversity of chalcidoid wasps in LUBout was found much more than LUBin. Biodiversity studies showed that the KA station has the greatest values of diversity, evenness, and richness of Chalcidoidea. This station is located outside of Lake Urmia basin with the warmest and most humid climate and forest ecosystem on the margins of the Aras River.
... Hymenoptera, one of four mega-diverse insect orders, comprises more than 155,000 described species [1,2] and plays a fundamental role in almost all terrestrial ecosystems [3,4]. Hymenoptera consists of the suborders of Apocrita and Symphyta [5]. ...
Three new species, Brachyoxyela conjunctiva sp. nov., Brachyoxyela leei sp. nov. and Brachyoxyela elliptica sp. nov., in the tribe Macroxyelini, subfamily Macroxyelinae of the family Xyelidae, are described and illustrated. All type specimens were collected from the Lower Cretaceous (Barremian–Aptian) Yixian Formation. Brachyoxyela conjunctiva sp. nov. and Brachyoxyela leei sp. nov. were from Huangbanjigou, Beipiao City, Liaoning Province, China, and Brachyoxyela elliptica sp. nov. from Sandaogou, Duolun County, Inner Mongolia, China. These specimens are assigned to a previously erected genus Brachyoxyela Gao, Zhao and Ren, 2011 based on the characters: vein Sc joining R distad origin of Rs and antenna with thread (terminal part of flagellum distal to the enlarged first flagellomere) longer than the first flagellomere. A key to species of Brachyoxyela is provided.
... The Braconidae is the second-largest parasitoid wasp family in Hymenoptera represent a cosmopolitan family having an estimated 40,000 species worldwide (Shaw and Huddleston, 1991;Goulet and Huber, 1993) and 19,411 are described under 1,062 genera and 46 subfamilies (Aguiar et al., 2013). Braconidae are primary parasitoids of other insects, with rare exceptions, and have high host specificity in each subfamily (Matthews, 1974;Marsh, 1979). ...
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The diversity and distribution of braconid wasps in the two biotic provinces of the North-East India biogeographic zone are provided. This compilation revealed 153 species under 66 genera belonging to 15 subfamilies.The maximum number of species have been recorded from Meghalaya with 97 species (63%), followed by Manipur with 32 (21%), Assam with 26 (17%)and Tripura with 16 (10%). Much work remains to be done on inventorying the species from these provinces; only 4 species under 2 subfamilies were recorded from Nagaland and one species from Mizoram. Out of 153 species, 51 species (33%) are endemic exclusively to this region.
... The parasitic Hymenoptera (ichneumonids, braconids, chalcids and platygastrids) are utilized in insect pest management. The order is divided into two suborders, Symphyta and Apocrita, it comprised of 27 superfamilies and 132 families (Aguiar et al., 2013 ...
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Altogether 260 species belonging to 82 genera under 17 families and 6 superfamilies are reported from North-East biogeographic zone of India. The maximum species diversity is reported from North-East Hills (216 species) followed by North East Brahmaputra Valley (59 species).
... Order HYMENOPTERA Hymenoptera is one of the megadiverse insect order having 2 suborders, 27 superfamilies, 116 families, 7,738 genera and 1,50,659 species (Aguiar et al., 2013) globally. The recent data states the presence of 19 superfamilies, 68 families and 12,605 species of Hymenoptera in India (Chandra, 2011). ...
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North-East Zone of India constitutes 5.2 per cent of total geographical area endowed with rich biodiversity. This zone depicts the faunistic and floristic resemblance of Indo-Malayan and Indo-Chinese bio-geographical regions. Zoological Survey of India made an attempt to compile all the available information on faunal diversity based on its surveys conducted over a period of more than a hundred years and the published literature elsewhere. It is revealed that altogether 18,527 species in 15 phyla of both Protozoa (243) and Animalia (18,284) have been documented, representing about 18% of the total Indian fauna (1,02,161 species). Phylum Arthropoda with 14,956 species represents 80.7%, of the overall faunal diversity of North-East India. With regards to faunal diversity in its biotic provinces and states, North-East Hills includes 14,838 species, Brahmaputra Valley 8,832 species, Meghalaya 9,853 species, Assam 9,607 species, Manipur 5,036 species, Nagaland 3,869 species, Tripura 3,605 species, and Mizoram 3,020 species. Out of 718 threatened species from India assessed by IUCN, 153 species are also distributed in North-East India, representing about 22.35% of total Indian threatened fauna. To protect the unique biodiversity of North-East Bio-geographical zone, 174 Protected Areas have been notified by Government of India.
... Hymenoptera, one of four mega-diverse insect orders, comprises more than 155,000 described species [1,2] and plays a fundamental role in almost all terrestrial ecosystems [3,4]. Hymenoptera consists of the suborders of Apocrita and Symphyta [5]. ...
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One new genus and species, Leptoxyela eximia gen. et sp. nov., and one new species, Scleroxyela cephalota sp. nov., are described and illustrated based on two well-preserved compression fossils from the Lower Cretaceous Yixian Formation and the Middle Jurassic Jiulongshan Formation of China, respectively. Leptoxyela eximia gen. et sp. nov. is placed in the tribe Angaridyelini, 1966 and Scleroxyela cephalota sp. nov. in the tribe Xyeleciini Benson, 1945; while both tribes are in the subfamily Macroxyelinae Ashmead, 1898 of Xyelidae. A key to the genera of Angaridyelini is provided. In addition, we investigated various angles between Rs+M and 1-Rs for known fossil species of Macroxyelinae, and we found the angle and the length of 1-Rs are correlated; however, we could not see any correlation between the angles and the fossil ages even within a tribe. Furthermore, based on Sc2 connected to R before Rs, the angle between Rs+M and 1-Rs, and the length of the first flagellomere, we believe that the tribe Ceroxyelini may have only one genus of Ceroxyela, and suggest that Isoxyela and Sinoxyela should be transferred to Gigantoxyelini.
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As a result of this study on proctotrupid wasps in Iran, 15 species belonging to seven genera were identified. Among them, seven species are new records for the Iranian fauna: Mischoserphus obesus Townes, 1981, Pschornia minora Townes, 1981, Exallonyx certus Townes, 1981, Exallonyx longicornis (Nees, 1834), Exallonyx confusus (Nixon, 1938), Exallonyx pallidistigma Morley, 1922 and Exallonyx subserratus Kieffer, 1908. The genus Pschornia Townes, 1981 is recorded from Iran for the first time and Pschornia persica Izadizadeh, Kolyada & Talebi sp. nov. is described as new for science. The number of species of Proctotrupidae in Iran is now raised from 18 to 26. Diagnostic characters of the newly recorded species are provided along with illustrations. A key to all genera and species of Proctotrupidae known in Iran, as well as a distribution map of all species throughout the world, is presented.
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The study included a survey of six geographical locations in Basra province, Southern Iraq, which are the Shatt al-Arab, Al-Hartha, Abu Al-Khasib, Al-Zubeyr, Al-Faw, and Qurna regions for the period from 1/2/2018 to 1/11/2019. To study the presence of wild bees, the results of the taxonomic study showed of samples collected for the diagnosis of 7 species or morphospecies, traced to four genera its (Ceratina , Andrena , Megachile and Icteranthidium ) and belonged to four tribes, three subfamily, and three families. It is less than was recorded in the old years.
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The study included a survey of six geographical locations in Basra province, Southern Iraq, which are the Shatt al-Arab, Al-Hartha, Abu Al-Khasib, Al-Zubeyr, Al-Faw, and Qurna regions for the period from 1/2/2018 to 1/11/2019. To study the presence of wild bees, the results of the taxonomic study showed of samples collected for the diagnosis of 7 species or morphospecies, traced to four genera its (Ceratina , Andrena , Megachile and Icteranthidium ) and belonged to four tribes, three subfamily, and three families. It is less than was recorded in the old years
The Stephanidae literature through July 2004 is summarized and 326 valid species, including 6 fossil species, are cataloged. Information on the localization of primary and secondary type material, as well as updated names for the localities of primary types, are provided for all valid and synonymized species. Literature treatment of male and female specimens is discriminated in the synonymic listing of all species. The following 90 nomenclatural changes are made: Genus-level type designation ( 1): Bothrioceros Europaeus Sichel designated as type species of Bothrioceros Sichel. New combinations ( 69): From Diastephanus: Foenatopus alutaceus (Morley), F. anupam (Narendran & Sureshan), F. bilineatus (Elliott), F. burmaensis ( Narendran & Sureshan), F. capitatus (Benoit), F. carinifrons (Enderlein), F. chinnarensis ( Sureshan), F. chinensis ( Elliott), F. christineae ( Narendran), F. costifrons ( Elliott), F. daccaensis ( Narendran & Sureshan), F. dohrni ( Enderlein), F. elegans ( Elliott), F. elegantulus ( Elliott), F. elongatus ( Elliott), F. equatorialis ( Benoit), F. flaviceps ( Elliott), F. flavifrons ( Elliott), F. frontilinea ( Morley), F. fuscidens (Kieffer), F. gracilis ( Kieffer), F. keralensis ( Narendran & Sureshan), F. lucifer ( Elliott), F. maculifemur ( Enderlein), F. multicolor ( Elliott), F. parviceps ( Enderlein), F. parvulus ( Elliott), F. priyae ( Narendran & Sureshan), F. quadridens (Elliott), F. salomonis (Westwood), F. sangalensis (Benoit), F. semiglaber (Elliott), F. simillimus (Elliott), F. stom (Narendran & Sureshan), F. sudhae (Narendran & Sureshan), F. sulcatus ( Elliott), F. szepligetii ( Enderlein), F. tertianus ( Morley), F. trialbatus ( Elliott), F. trilineatus ( Elliott), F. trilobatus ( Elliott), F. wynadensis ( Sureshan Narendran); from Madegafoenus: Megischus bekilyanus ( Benoit), M. occiputalis ( Benoit); from Megischus: Afromegischus gigas (Schletterer), Foenatopus phoberopus (Sausurre); from Neostephanus: Foenatopus alluaudi ( Kieffer), F. berlandi ( Benoit), F. camerunus ( Enderlein), F. collaris ( Benoit), F. crassiceps (Bischoff), F. globiceps ( Enderlein), F. insignis ( Schletterer), F. longicaudatus ( Benoit), F. micans ( Benoit), F. obockensis ( Benoit), F. oemidaphagus ( Benoit), F. pauliani ( Benoit), F. pentheri ( Kieffer); from Pseudomegischus: Afromegischus tibiator ( Schletterer); from Stephanus: Comnatopus xanthocephalus ( Cameron); Megischus collectivus ( Elliott), M. diversus ( Schletterer), M. lanceolatus ( Kieffer), M. sanmartinianus ( Orfila), M. seyrigi ( Benoit), M. hornianus ( Enderlein), M. tricolor ( Elliott), M. willineri ( Orfila). Replacement names ( 3): Foenatopus annularis Aguiar, generic transfer and replacement name for Diastephanus annulipes Elliott, preventing secondary homonym with F. annulipes ( Elliott); F. rufocinctus Aguiar, new status and replacement name for Stephanus togoensis var. fasciatus Enderlein, preventing secondary homonym with F. fasciatus Szepligeti; Parastephanellus curtus Aguiar, replacement name for secondary homonym of P. curticollis ( Elliott) with P. curticollis Elliott. New status ( 2): Comnatopus Achterberg, new status for Profoenatopus ( Comnatopus) Achterberg; Parastephanellus coriaceus Kieffer, new status for Parastephanellus polychromus var. coriacea Kieffer. New genus-group synonyms ( 1): Pseudomegischus ( Callomegischus) Achterberg, new synonymy for Afromegischus Achterberg. New species-group synonyms ( 7): Stephanus comma Morley, new synonymy for Comnatopus xanthocephalus ( Cameron); Foenatopus flavicollis flavissimus Benoit, new synonymy for F. flavicollis Cameron; Stephanus testaceipes Elliott, Stephanus iesuiticus Orfila, S. paraguayensis Orfila, and S. martinezi Orfila, new synonyms for Megischus maculipennis Westwood; Parastephanellus impunctatus Elliott, new synonymy for P. nigricaudus ( Sichel). Reinstated name ( 1): Megischus ruficeps Saussure, reinstated, invalidating M. saussurei (Schulz). Species inquirendum ( 1): Schlettererius rufipes ( Say, 1824), a Braconidae. Nomina nuda ( 3): Foenus diadema Fabricius, nomen nudum for M. coronator ( Fabricius); Stephanus curticauda Elliott, nomen nudum for M. curtus ( Elliott); Stephanus frontalis Klug, nomen nudum, correspondent species unknown.
The Mexican genera and species of Stephanidae, and the western species of Foenatopus Smith are revised. Twelve species are recognized, two in Foenatopus and ten in Megischus Brulle, all but one occurring in Mexico. Six new species of Megischus are described, M. anaxeus sp. n., M. celaenocephalus sp. n., M. jaliscoensis sp. n., M. melanogladius sp. n., M. mexicanus sp. n., and M. peninsularis sp. n. One genus ( Foenatopus Smith) and four species are registered for the first time in Mexico, F. annulipes Kieffer, M. arizonicus Townes, M. bicolor ( Westwood), and M. niger Smith. Many new distribution records in Mexico and intensive new morphological variation are registered, illustrated and discussed for M. texanus Townes. Much new morphological variation is reported and illustrated for M. arizonicus. The male of F. aurantiiceps Brues is reported for the first time. Mexico is identified as an area of overlapping distribution between the essentially North American M. bicolor and the typically Central American M. niger; intermediate forms between these two species are described, and new diagnostic features are proposed for their identification. Some of the treated species are proposed as possibly highly endemic, occurring only in particular regions in the country, namely M. peninsularis, restricted to Baja California, and M. celaenocephalus, restricted to central Mexico. Some of the morphological variation observed for F. aurantiiceps is tentatively linked to specimens occurring in the south vs. north of the Amazon river, and to localities of high vs. low altitude. General stephanid morphology is illustrated, its correspondent terminology updated, and a key is provided for recognition of all treated genera and species. Full description is provided for each new taxon, and photographic illustrations are provided for all valid taxa. Extensive biometric data, and RGB formulas for color variation are provided for nearly all treated species.
Protobethylus eocenicus, new genus and species of Bethylidae: Bethylinae, is described from the Lowermost Eocene amber of the Paris Basin (France). A new phylogenetic analysis of the bethyline genera is proposed. Although having superficial similarities with the habitus of Lytopsenella and Eupsenella; Protobethylus is more derived than these basal genera.
Dryinus alamellatus Olmi & Guglielmino, sp. nov. is described from amber collected in Dominican Republic. A revision and a key to the fossil Neotropical species of Dryinus Latreille 1804 belonging to constans group is presented.