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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)

Authors:

Abstract

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 http://creativecommons.org/licenses/by/3.0
51
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334 (online edition)
Copyright © 2013 Magnolia Press
Zootaxa 3703 (1): 051062
www.mapress.com
/
zoota xa
/
Correspondence
http://dx.doi.org/10.11646/zootaxa.3703.1.12
http://zoobank.org/urn:lsid:zoobank.org:pub:AB0CBAED-2CC8-4338-B6C3-F93C0E437C70
Order Hymenoptera*
ALEXANDRE P. AGUIAR
1
, ANDREW R. DEANS
2
, MICHAEL S. ENGEL
3
, MATTIAS FORSHAGE
4
,
JOHN T. HUBER
5
, JOHN T. JENNINGS
6
, NORMAN F. JOHNSON
7
, ARKADY S. LELEJ
8
, JOHN T.
LONGINO
9
, VOLKER LOHRMANN
10
, ISTVÁN MI
11
, MICHAEL OHL
12
, CLAUS RASMUSSEN
13,**
,
ANDREAS TAEGER
14
& DICKY SICK KI YU
15
1
Universidade Federal do Espírito Santo, Depto. de Ciências Biológicas, Av. Fernando Ferrari, s/n – Goiabeiras, 29075-010 Vitoria,
ES, BRAZIL, e-mail:
aguiar.2@osu.edu
2
Department of Entomology, Pennsylvania State University, 501 ASI Building, University Park, PA 16802, USA,
e-mail: adeans@psu.edu
3
Division of Entomology, Natural History Museum, 1501 Crestline Drive – Suite 140, University of Kansas, Lawrence, KS 66045,
USA,
e-mail: msengel@ku.edu
4
Swedish Museum of Natural History, Department of Entomology, Box 50007, SE-104 05 Stockholm, SWEDEN,
e-mail: mattias.forshage@nrm.se
5
Canadian National Collection of Insects (CNC), Agriculture and Agri-Food Canada, 960 Carling Avenue, Ottawa, ON, K1A 0C6,
CANADA,
e-mail: john.huber@agr.gc.ca
6
School of Earth & Environmental Sciences, G15d Darling Building, The University of Adelaide, 5005, AUSTRALIA,
e-mail: john.jennings@adelaide.edu.au
7
Department of Evolution, Ecology & Organismal Biology, Director C.A. Triplehorn Insect Collection, The Ohio State University,
1315 Kinnear Road, Columbus, OH 43212, USA,
e-mail: johnson.2@osu.edu
8
Institute of Biology and Soil Science, Russian Academy of Sciences, Vladivostok-22, 690022, RUSSIA, e-mail: lelej@biosoil.ru
9
Department of Biology, 257 S 1400 E, University of Utah, Salt Lake City, UT 84112, USA, e-mail: jacklongino@gmail.com
10
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,
e-mail: Volker.Lohrmann@mfn-berlin.de
11
Department of Entomology, Pennsylvania State University, 501 ASI Building, University Park, PA 16802, USA,
e-mail: istvan.miko@gmail.com
12
Museum für Naturkunde, Leibniz-Institut für Evolutions- und Biodiversitätsforschung, Invalidenstraße 43, 10115 Berlin, GERMANY,
e-mail: Michael.Ohl@mfn-berlin.de
13
Department of Bioscience, Aarhus University, Ny Munkegade 114, Bldg. 1540, DK-8000 Aarhus C, DENMARK,
e-mail: alrunen@yahoo.com
14
Senckenberg Deutsches Entomologisches Institut (SDEI), Eberswalder Str. 90, 15374 Müncheberg, GERMANY,
e-mail: Andreas.Taeger@senckenberg.de
15
403-1220 Merivale Road, Ottawa, Ontario K1Z 8P2, CANADA, e-mail: dickyyu@gmail.com
** 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.
Abstract
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
AGUIAR ET AL.: HYMENOPTERA
52 · Zootaxa 3703 (1) © 2013 Magnolia Press
Introduction
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,
1984).
Classification
Order Hymenoptera Linnaeus, 1758 (2 suborders) (8,423 (and †684) genera, 152,677 (and †2,428) species)
Suborder Symphyta
1
(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
ZHANG (ED.): ANIMAL BIODIVERSITY (ADDENDA 2013)
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
2
Handlirsch, 1906 (1 family)
† Family Ephialtitidae Handlirsch, 1906 (†2 genera, †60 species)
Superfamily Trigonaloidea
3
Cresson, 1887 (2 families)
† Family Maimetshidae
4
Rasnitsyn, 1975 (†9 genera, †11 species)
Family Trigonalidae
5
Cresson, 1887 (16 (and †4) genera, 92 (and †5) species)
Superfamily Megalyroidea Schletterer, 1889 (1 family)
Family Megalyridae
6
Schletterer, 1889 (8 (and †11) genera, 43 (and †40) species)
Superfamily Stephanoidea Leach, 1815 (1 family)
Family Stephanidae
7
Leach, 1815 (11 (and †4) genera, 342 (and †8) species)
Superfamily Ceraphronoidea Haliday, 1833 (4 families)
Family Ceraphronidae
8
Haliday, 1833 (15 genera, 304 (and †2) species)
Family Megaspilidae
9
Ashmead, 1893 (12 (and †1) genera, 299 (and †12) species)
2. http://www.taxapad.com/taxapadmain.php.
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
Trigonalidae.
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.
AGUIAR ET AL.: HYMENOPTERA
54 · Zootaxa 3703 (1) © 2013 Magnolia Press
† Family Radiophronidae
10
Ortega-Blanco, Rasnitsyn & Declòs, 2010 (†2 genera, †2 species)
† Family Stigmaphronidae
11
Kozlov, 1975 (†8 genera, †25 species)
Superfamily Evanioidea Latreille, 1802 (5 families)
† Family Andreneliidae
12
Rasnitsyn & Martínez-Declòs, 2000 (†1 genus, †1 species)
Family Aulacidae
13
Shuckard, 1841 (2 (and †10) genera, 185 (and †18) species)
Family Evaniidae
14
Latreille, 1802 (21 (and †9) genera, 449 (and †19) species)
Family Gasteruptiidae
15
Ashmead, 1900 (6 genera, 496 species)
† Family Praeaulacidae
16
Rasnitsyn, 1972 (†10 genera, †39 species)
Superfamily Mymarommatoidea Debauche, 1948 (3 families)
† Family Alavarommatidae
17
Ortega-Blanco, Peñalver, Delclòs & Engel, 2011 (†1 genus, †1
species)
† Family Gallorommatidae
18
Gibson, Read & Huber, 2007 (†1 genus, †3 species)
Family Mymarommatidae
19
Debauche, 1948 (3 (and †2) genera, 10 (and †10) species)
Superfamily Proctotrupoidea Latreille, 1802 (11 families)
Family Austroniidae
20
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
21
Kozlov, 1970 (†16 genera, †33 species)
Family Pelecinidae
22
Haliday, 1839 (1 (and †5) genera, 3 (and †8) species)
Family Peradeniidae
23
Naumann & Masner, 1985 (1 genus, 2 (and †1) species)
Family Proctorenyxidae
24
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
25
Haliday, 1833 (190 (and †7) genera, 2048 (and †22) species)
Family Ismaridae
26
Thomson, 1858 (1 genus, 29 species)
Family Maamingidae
27
Early, Masner, Naumann & Austin, 2001 (1 genus, 2 species)
Family Monomachidae
28
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.
21. http://osuc.biosci.ohio-state.edu.
22. Engel & Grimaldi 2006a.
23. Johnson et al. 2001.
24. Lelej & Kozlov 1999; He et al. 2002.
25. Perrichot & Nel 2008.
26. http://osuc.biosci.ohio-state.edu.
27. Early et al. 2001.
28. Johnson & Musetti 2012.
Zootaxa 3703 (1) © 2013 Magnolia Press · 55
ZHANG (ED.): ANIMAL BIODIVERSITY (ADDENDA 2013)
† Family Spathiopterygidae
29
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
30
Brues, 1937 (†3 genera, †9 species)
Superfamily Cynipoidea Latreille, 1802 (8 families)
Family Austrocynipidae
31
Riek, 1971 (1 genus, 1 species)
Family Cynipidae
32
Latreille, 1802 (74 (and †3) genera, 1412 (and †11) species)
Family Figitidae Hartig, 1840 (148 (and †10) genera, 1571 (and †11) species)
† Family Gerocynipidae
33
Liu & Engel, 2007 (†3 genera, †5 species)
Family Ibaliidae
34
Thomson, 1862 (3 (and †1) genera, 20 (and †2) species)
Family Liopteridae Ashmead, 1895 (10 (and †2) genera, 153 (and †2) species)
† Family Protimaspidae
35
Liu & Engel, 2007 (†1 genus, †1 species)
† Family Stolamissidae
36
Liu & Engel, 2007 (†1 genus, †1 species)
Superfamily Chalcidoidea
37
Latreille, 1817 (23 families)
Family Agaonidae
38
Walker, 1846 (20 (and †2) genera, 762 (and †4) species)
Family Aphelinidae Thomson, 1876 (29 (and †1) genera, 1078 (and †1) species)
Family Azotidae
39
Nikol’skaya & Yasnosh, 1966 (1 genus, 92 species)
Family Chalcididae Latreille, 1817 (90 (and †3) genera, 1469 (and †5) species)
Family Cynipencyrtidae
40
Trjapitzin, 1973 (1 genus, 1 species)
Family Encyrtidae Walker, 1837 (493 (and †2) genera, 4058 (and †3) species)
Family Eriaporidae
41
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
42
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
43
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.
AGUIAR ET AL.: HYMENOPTERA
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
44
Latreille, 1802 (1575 genera, 24025 (and †216) species)
† Family Praeichneumonidae
45
Rasnitsyn, 1983 (†1 genus, †5 species)
Superfamily Bethylonymoidea Rasnitsyn, 1975 (1 family)
† Family Bethylonymidae
46
Rasnitsyn, 1975 (†2 genera, †17 species)
Superfamily Chrysidoidea Latreille, 1802 (9 families)
Family Bethylidae
47
Haliday, 1839 (84 (and †11) genera, 2340 (and †13) species)
Family Chrysididae
48
Latreille, 1802 (81 (and †6) genera, 2500 (and †9) species)
Family Dryinidae
49
Haliday, 1833 (41 (and †2) genera, 1605 (and †19) species)
Family Embolemidae
50
Förster, 1856 (2 (and †1) genera, 39 (and †9) species)
† Family Falsiformicidae
51
Rasnitsyn, 1975 (†1 genus, †1 species)
† Family Plumalexiidae
52
Brothers, 2011 (†1 genus, †1 species)
Family Plumariidae
53
Bischoff, 1914 (7 genera, 22 species)
Family Sclerogibbidae
54
Ashmead, 1902 (3 (and †1) genera, 20 (and †2) species)
Family Scolebythidae
55
Evans, 1963 (4 (and †5) genera, 6 (and †7) species)
Superfamily Vespoidea
56
Latreille, 1802 (10 families)
Family Bradynobaenidae
57
de Saussure, 1892 (10 genera, 188 species)
Family Formicidae
58
Latreille, 1802 (299 (and †132) genera, 12199 (and †620) species)
Family Mutillidae
59
Latreille, 1802 (210 (and †1) genera, 4302 (and †12) species)
Family Pompilidae
60
Latreille, 1804 (125 (and †2) genera, 4855 (and †16) species)
Family Rhopalosomatidae
61
Ashmead, 1896 (4 (and †4) genera, 72 (and †4) species)
Family Sapygidae
62
Latreille, 1810 (12 (and †1) genera, 66 (and †1) species)
Family Scoliidae
63
64
Latreille, 1802 (143 (and †5) genera, 560 (and †17) species)
Family Sierolomorphidae
65
Krombein, 1951 (2 genera, 11 (and †1) species)
Family Tiphiidae
66
Leach, 1815 (120 (and †5) genera, 2000 (and †17) species)
Family Vespidae
67
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. http://antcat.org (accessed March 2012).
59. Brothers 2003; Lelej 2005.
60. Engel & Grimaldi 2006c.
61. http://rhopalosomatidae.hymis.eu (accessed August 2013).
62. Kurzenko 2009; Bennett & Engel 2005.
Zootaxa 3703 (1) © 2013 Magnolia Press · 57
ZHANG (ED.): ANIMAL BIODIVERSITY (ADDENDA 2013)
Superfamily Apoidea Latreille, 1802 (13 families)
Family Ampulicidae
68
Shuckard, 1840 (6 (and †7) genera, 200 (and †8) species)
Family Andrenidae
69
Latreille, 1802 (77 (and †3) genera, 2917 (and †11) species)
† Family Angarosphecidae
70
Rasnitsyn, 1975 (†13 genera, †44 species)
Family Apidae
71
Latreille, 1802 (209 (and †22) genera, 5749 (and †87) species)
Family Colletidae
72
Lepeletier de Saint Fargeau, 1841 (86 genera, 2547 (and †2) species)
Family Crabronidae
73
Latreille, 1802 (242 (and †24) genera, 8773
74
(and †29) species)
Family Halictidae
75
Thomson, 1869 (79 (and †7) genera, 4327 (and †22) species)
Family Megachilidae
76
Latreille, 1802 (76 (and †6) genera, 4096 (and †34) species)
Family Melittidae
77
Schenck, 1860 (15 (and †2) genera, 187 (and †4) species)
† Family Mellitosphecidae
78
Poinar & Danforth, 2006 (†1 genus, †1 species)
† Family Paleomelittidae
79
Engel, 2001 (†1 genus, †1 species)
Family Sphecidae
80
Latreille, 1802 (19 (and †1) genera, 724
81
(and †1) species)
Family Stenotritidae
82
Cockerell, 1934 (2 genera, 21 species)
Apocrita incertae sedis (3 families)
† Family Archaeocynipidae
83
Rasnitsyn & Kovalev, 1988 (†2 genera, †3 species)
† Family Eostephanitidae Hong, 2002 (†1 genus, †1 species)
† Family Kuafuidae
84
Rasnitsyn & Zhang, 2010 (†3 genera, †3 species)
Acknowledgements
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.
68. http://research.calacademy.org/ent/catalog_sphecidae.
69. Michener 2007; Michez et al. 2012; Ascher & Pickering 2012.
70. http://research.calacademy.org/ent/catalog_sphecidae.
71. Michener 2007; Michez et al. 2012; Ascher & Pickering 2012.
72. Michener 2007; Michez et al. 2012; Ascher & Pickering 2012.
73. http://research.calacademy.org/ent/catalog_sphecidae.
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.
80. http://research.calacademy.org/ent/catalog_sphecidae.
81. Updated August 2013.
82. Michener 2007; Ascher & Pickering 2012.
AGUIAR ET AL.: HYMENOPTERA
58 · Zootaxa 3703 (1) © 2013 Magnolia Press
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... 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. ...
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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.
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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.
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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.