ISSN 1911-2173

Key to the New World genera of Braconinae (Hymenoptera: Ichneumonoidea: Braconidae) and synopses of the genera

Accepted: September 24, 2024

Donald L. J. Quicke1*

Buntika A. Butcher1

Michael Sharkey2

1Integrative Insect Ecology Research Unit, Department of Biology, Faculty of Life Sciences, Chulalongkorn University, Bangkok, Thailand

2Hymenoptera Institute, 1339 La Loma Dr., California, 92373, USA

*Corresponding author: [email protected]

Key to the New World genera of Braconinae (Hymenoptera: Ichneumonoidea: Braconidae) and synopses of the genera

Key to the New World genera of Braconinae (Hymenoptera: Ichneumonoidea: Braconidae) and synopses of the genera

Accepted: September 24, 2024

Donald L. J. Quicke1*

Buntika A. Butcher1

Michael Sharkey2

1Integrative Insect Ecology Research Unit, Department of Biology, Faculty of Life Sciences, Chulalongkorn University, Bangkok, Thailand

2Hymenoptera Institute, 1339 La Loma Dr., California, 92373, USA

*Corresponding author: [email protected]

Abstract

An updated key is provided to the 31 genera of Braconinae found in the New World (Western Hemisphere), including Greenland and the Caribbean Islands. It is based upon the keys by Quicke (1997) and Sharkey et al. (2021a). The key is fully illustrated and is meant to facilitate use by non-specialists; the online version additionally follows web accessibility standards and is usable on desktop and mobile devices. An overview of the subfamily, and one or two pages dedicated to each genus with further extensive illustrations are included.

Bracon sp. from Mompha epilobiella (Denis and Schiffermüller) Fergus, Ontario, Canada, on Epilobium parviflorum (Schreb.). Photo taken by Steve Marshall.
Bracon sp. from Mompha epilobiella (Denis and Schiffermüller) Fergus, Ontario, Canada, on Epilobium parviflorum (Schreb.). Photo taken by Steve Marshall.

Quicke, D.L.J., 1997. Subfamily Braconinae, pp.149-174. In: Wharton, R.A., Marsh, P.M., & Sharkey, M.J. (eds). Manual of the New World genera of Braconidae (Hymenoptera). Special Publication of the International Society of Hymenopterists, 439 pp.

Sharkey, M.J., Janzen, D.H., Hallwachs, W., Chapman, E.G., Smith, M.A., Dapkey, T., Brown, A., Ratnasingham, S., Naik, S., Manjunath, R., Perez, K., Milton, M., Hebert, P., Shaw, S.R., Kittel, R.N., Solis, M.A., Metz, M.A., Goldstein, P.Z., Brown, J.W., Quicke, D.L.J., van Achterberg, C., Brown, B.V. and Burns, J.M. 2021a. Minimalist revision and description of 403 new species in 11 subfamilies of Costa Rican braconid parasitoid wasps, including host records for 219 species. ZooKeys, 1013: 1-665. https://doi.org/10.3897/zookeys.1013.55600

Introduction

The Braconinae is a large subfamily of moderately small to large wasps with more than 2000 described species and almost 200 genera worldwide (Shenefelt 1978; Shaw and Huddleston 1991; Quicke et al. 2023). They are most diverse in the Old World tropics but are also well represented in the New World. Comparatively few studies have been made on the New World fauna. Many New World species are erroneously included under the generic names Iphiaulax Förster (a genus that has an entirely Old World distribution), Ipobracon Thomson (a junior synonym of Cyanopterus Haliday), or Bracon Fabricius. Iphiaulax and Bracon (=Microbracon Ashmead) have been used in a far wider sense than they are today. Many of these species have subsequently been reclassified by Quicke (1988a, 1989a, 1991) and Quicke and Genise (1994), mostly moving them into Digonogastra Ashmead. The aberrant Neotropical genus Gnathobracon Costa was redescribed and transferred formally from Rogadinae to Braconinae (Quicke and Huddleston 1991). Several new genera have been described over the past 30 or so years by Shenefelt (in Mason 1978), Quicke (1988b, 1989b, 1994, 1995), van Achterberg (1989), Quicke and Sharkey (1989), Quicke and Delobel (1995), and Quicke et al. (2020b).

Overview of subfamily

Phylogeny

Belshaw et al. (2001) published a phylogenetic analysis based on a limited number of genera with a combined 28S rDNA and morphological data set. Quicke et al. (2023) conducted a molecular-based phylogenetic analysis of the subfamily employing the following markers: mitochondrial cytochrome c oxidase subunit I, 16S rDNA, nuclear 28S D2-D3 rDNA and elongation factor 1-alpha. They revised the tribal classification, placed most genera to tribe and synonymized Alienoclypeus with Atanycolus. In that paper they erected a new, tribe, Compsobraconii to include a small number of entirely New World taxa that were recovered as a fairly basal clade within the subfamily.

Biology

With a very few exceptions, New World braconines are idiobiont ectoparasitoids of concealed holometabolous insect larvae, especially of  Lepidoptera and Coleoptera, though a few species, mostly in the genus Bracon, attack concealed larvae of Diptera or 'Symphyta' (Hymenoptera). This is true for the whole subfamily except for the small, entirely Old World tropical subtribe Aspidobraconina whose included species are endoparasitoids of butterfly pupae, one known entomophytophagous species from India (Ranjith et al. 2016), and one completely predatory species (on a gall-forming mite) (Ranjith et al. 2022), also from India. Braconines are synovigenic and their large eggs are usually laid on a host that was previously paralyzed by injection of venom (Shaw and Huddleston 1991). Host feeding by adults appears to be quite common among species of Bracon and Habrobracon and sometimes involves the construction of feeding tubes (Schöller 2010). Both solitary and gregarious parasitism occur, sometimes with closely related species displaying different strategies (Wharton et al. 1989). Some genera appear to be somewhat specialized in their host range though most species are at least oligophagous. For example, the species of Myosoma Brullé, are parasitoids of stem-boring Lepidoptera (Pyralidae and Sesiidae), while species of Coeloides Wesmael are specialized on bark-boring Coleoptera larvae. The biologies of many Neotropical genera are unknown but members of one recently described genus, Cyclaulacidea Quicke and Delobel, have been reared from Bruchinae (Chrysomelidae) larvae, and most are presumed to be typical idiobiont ectoparasitoids. More aberrant biologies include one Compsobraconoides species that is an ectoparasitoid of adult Azteca ants (Yu and Quicke 1997), and a species group of Bracon that is a phytophagous seed-predator (Flores et al. 2006; Perioto et al. 2011).

Several species have been used in biocontrol programs in the New World. Various species of Bracon, Habrobracon Ashmead and Bracon (Pigeria) van Achterberg have been released to control a range of insects including stored product and stem borer pests. For example, Habrobracon brevicornis (Wesmael) has been released on numerous occasions in the USA against Heliothis virescens (F.), Helicoverpa zea (Boddie), Ostrinia nubilalis (Hubner) and Pectinophora gossypiella (Saunders), while Bracon (Pigeria) piger (Wesmael) has been released against Etiella zinckenella (Treit) and Cnephasia spp. Bracon mellitor Say is one of the most important natural parasitoids of the boll weevil, Anthonomus grandis Boheman, in the USA (Tillman and Gate 1989) while various other braconines are important in the natural regulation of bark beetle pests in North American forests. Members of several endemic New World genera are at least locally important parasitoids of pyralid cane borers, Diatraea spp., in Central and South America, including Digonogastra kimballi Kirkland, Palabracon diairaeaphilus Quicke, and Myosomatoides myersi Quicke (= pennipes Myers).

Diversity

There are probably tens of thousands of species of Braconinae worldwide but only about 3,000 species have been described (Yu et al. 2016). Species of the majority of New World genera are only infrequently collected but those of two species-rich genera, Bracon and the generally larger and often brightly colored Digonogastra, are abundant almost everywhere. Presently (Jan. 20, 2024), there are 1342 BINs (barcode index numbers, proxies for species (Rasnasingham and Hebert 2013) on BOLD, the Barcode of Life database, http://www.boldsystems.org); this includes 206 from Canada and 303 from Costa Rica, two of the best-collected countries in the World. Many more are to be found in both countries, especially Costa Rica.

Aguirre et al. (2017) found that braconine species had relatively narrow latitudinal ranges compared with those of the Doryctinae, which have broadly similar biology. Also, they showed that braconine species richness had two peaks in relation to altitude, one at 200m a.s.l., the second at approximately 1400m.

Distribution

The Braconinae have a cosmopolitan distribution, with the greatest generic, and probably specific, diversity being in the Old World tropics. The Nearctic fauna is similar in its generic composition to that of the Palaearctic, except for numerous species of Digonogastra which appears to be absent in the Palaearctic. The Neotropical genera are mostly endemic, except for Bracon, Vipio, and Digonogastra.

Distinguishing features

See Figs. 5-9 in Sharkey et al. (2023) for illustrations of morphological features. All New World species are distinctly cyclostome (i.e., have a large, dorsally rounded depression above the mandibles formed by a ventrally recessed clypeus and an exposed, concave and usually glabrous labrum), though in Gnathobracon and a few species in other genera the lower face and clypeal configuration are somewhat modified. The occipital carina is always absent (except in one aberrant Old World genus). The precoxal groove is almost always absent. Tergum 1 is usually distinctly flattened laterally with a domed apicomedial area and a midbasal depression. The hind wing cross vein m-cu is always absent and the hind wing vein M+Cu is less than 0.5 times the length of vein 1M (the latter nearly always being markedly thicker basally than distally). Most, if not all, Braconinae produce a distinctive, earthy-oily, smell when handled while alive and this can sometimes be noticed when walking through woodlands indicating the presence of larger species nearby.

Note. All images are by Sharkey unless otherwise noted and all imaged specimens are in the collection of the Hymenoptera Institute, unless otherwise noted.

Checklist

Table 1. List of the genera of Braconinae recognized in the New World.

 Acgorium Sharkey & Quicke, 2020

 Amyosoma Viereck, 1913

 Atanycolus Foerster, 1862

 Bracon Fabricius, 1804

 Calobracon Szépligeti, 1902

 Cervellus Szépligeti, 1904

 Chartobracon van Achterberg, 1983

 Coeloides Wesmael, 1838

 Compsobracon Ashmead, 1900

 Compsobraconoides Quicke, 1989

 Cyanopterus Haliday, 1835

 Cyclaulacidea Quicke and Delobel, 1995

 Cyclaulax Cameron, 1911

 Digonogastra Viereck, 1912

 Gnathobracon Costa, 1864

 Gozmanycomp Papp, 2007

 Gracilibracon Quicke, 1995

 Habrobracon Ashmead, 1895

 Hemibracon Szépligeti, 1906

 Lapicida Quicke, 1989

 Lasiophorus Haliday, 1838

 Leptobracon Szépligeti, 1901

 Megabracon Szépligeti, 1906

 Megacoeloides Quicke, 1989

 Myosoma Brullé, 1846

 Myosomatoides Quicke, 1994

 Palabracon Quicke, 1988

 Pheloura van Achterberg, 1989

 Sacirema Quicke, 1995

 Vipio Latreille, 1804

 Vipiomorpha Tobias, 1962

Key to genera

Key to the New World genera of Braconinae
Modified from Sharkey et al. (2021). Clicking the hyperlinked text in the key below will take the reader to the corresponding couplet or genus synopsis.
The shape of the antennal scape is important for generic identification. For determination of the relative lengths of the dorsal and ventral sides (couplets 4, 12, and 16), the antenna should be positioned as in couplet 12. The terms rare and uncommon are used conservatively, for example, the authors may never have encountered a rare genus outside of museum collections.
1.
A. Face with median projection that is approximately nearly round in cross section, although its apex sometimes bifurcate; Neotropical.

B. Face with large, horizontal, dorsally concave plate-like projection; Neotropical.

C. Face without a projection; widespread.

Figure for couplet 1

2.
A. Flagellomere 1 with thorn-like, pre-apical projection.

B. Flagellomere 1 simple or occasionally apically flared.

Figure for couplet 2

3(1).
A. Flagellomere 1, and sometimes 2 and 3, expanded apico-ventrally.

B. Flagellomere 1 simple, not expanded apico-ventrally.

Figure for couplet 3

4(3).
A. Scape shorter ventrally than dorsally (when antenna directed anteriorly); flagellum uniformly dark, without white or cream-colored band at mid-length; tergum 2 usually with sculpture; Nearctic and Mexico, common.

B. Scape longer ventrally than dorsally (antenna directed anteriorly); flagellum with white or cream-colored band at mid-length; tergum 2 entirely smooth; Neotropical, uncommon.

Figure for couplet 4

5(3).
A. Scape with false sub-apical margin, well-separated from the real margin, and scape abruptly and concavely narrowing towards base.

B. Scape without false sub-apical margin, and scape gradually narrowing towards base.

Figure for couplet 5

6(5).
A. Antennal socket greatly protruding anteriorly. Scape as wide as long. Temple protruding lateral to eye, i.e. wider than head across eyes; Neotropical, uncommon.

B. Antennal sockets slightly protruding anteriorly. Scape as long as wide. Temple shorter than eye.

Figure for couplet 6

7(6).
A. Tergum 2 with triangular or (rarely) quadrate raised area that narrows posteriorly; widespread.

B. Tergum 2 smooth and flat or with a smooth posteriorly widening “pinched-up” area; Neotropical.

Figure for couplet 7

8(7).
A. Raised area of tergum 1 with semi-circular emargination posterolaterally; Neotropical.

B. Raised area of tergum 1 without abrupt emargination posterolaterally; widespread.

Figure for couplet 8

9(7).
A. Metasomal tergum 2 strongly ‘pinched-up’ anteromedially.

B. Metasomal tergum 2 not or hardly ‘pinched-up’ anteromedially.
Cyclaulax (in part)

Figure for couplet 9

10(5).
A. Middle part of face clearly demarked from lateral parts by a pair of sub-medial, longitudinal carinae extending ventrally from antennal sockets. AA. Metasomal tergum 2 without mediobasal, posteriorly narrowing triangular area.

B. Middle part of face not clearly demarked from lateral parts by longitudinal grooves or carinae. BB. Metasomal tergum 2 often with mediobasal, posteriorly narrowing triangular area.

Figure for couplet 10

11(10).
A. Hypopygium truncate in lateral view.

B. Hypopygium acutely pointed in lateral view.

Figure for couplet 11

12(11).
A. Scape shorter ventrally (V) than dorsally (D) (when antenna directed anteriorly).

B. Scape longer ventrally (V) than dorsally (D) (when antenna directed anteriorly).

Figure for couplet 12

13(12).
A. Propodeum entirely smooth. Tergum 2 flat or weakly pinched anteromedially.

B. Propodeum with distinct longitudinal carina midposteriorly. Tergum 2 more pinched-up midanteriorly, especially in females.

Figure for couplet 13

14(12).
A. Length of tergum 2 more than half as long as tergum 3.

B. Length of tergum 2 less than half as long as tergum 3.
Cyclaulax (in part)

Figure for couplet 14

15(14).
A. Face rugose with a lot of granulate sculpture, often with a raised granulate area medially. AA. Raised median area of tergum 1 rather square in cross-section, its lateral margin more or less carinate posteriorly.

B. Face coarsely carinate-rugose medially with little or no granulate sculpture and without a raised granulate area medially. BB. Raised median area of tergum 1 more rounded in cross-section.

Figure for couplet 15

16(10).
A. Scape apicoIaterally not, or only weakly, emarginate. Scape shorter ventrally than dorsally (when antenna directed anteriorly).

B. Scape apicolaterally emarginate. Scape longer ventrally than dorsally (when antenna directed anteriorly).

Figure for couplet 16

17(16).
A. Marginal cell of forewing short, vein 3RSb reaching wing margin at most 0.7 times distance between apex of stigma and wing tip. AA. Clypeus with widely spaced pair or cluster of long setae that often touch apically and/or, AAA. Tergum 4 with characteristic pattern of fine striae that curve away from midline.

B. Marginal cell of forewing usually longer, vein 3RSb reaching wing margin at least 0.8 times distance between apex of stigma and wing tip; if shorter, then… BB. Clypeus without widely spaced pair or cluster long setae (clypeal guard setae evenly spaced and not touching apically). BBB. Tergum 4 without fine striae that curve away from the midline.

Figure for couplet 17

18(17).
A. Hind tibia strongly laterally compressed, very broad in profile with especially long setae. AA. Metasomal tergum 2 with midlongitudinal ridge; Neotropical, rare.

B. Hind tibia usually not strongly laterally compressed and usually without especially long marginal setae. BB. If rather strongly compressed with long marginal setae (most Myosoma species) then metasomal tergum 2 without midlongitudinal ridge; widespread, common.

Figure for couplet 18

19(18).
A. Body very strongly dorsoventrally depressed, mesosoma more than 2.8 times as long as maximum height; Holarctic, rare.

B. Body not or only moderately dorsoventrally depressed, mesosoma less than 2.3 times as long as maximum height; widespread, common.

Figure for couplet 19

20(19).
A. Median tergite 1 very long and narrow, more than 2.0 times as long as maximal width. AA. Median tergite 1 usually (in dead specimens) almost vertical and abutting propodeum, and therefore difficult to see.

B. Median tergum 1 not especially elongate, less than 2.0 times as long as maximum width.

Figure for couplet 20

21(20).
A. Hind femur and tibia with short pale setae; Neotropical.

B. Hind femur and hind tibia with long black setae; widespread.

Figure for couplet 21

22(20).
A. Propodeum with a complete midlongitudinal carina; rare.

B. Propodeum entirely smooth, or at most with a short midlongitudinal carina posteriorly; very common.

Figure for couplet 22

23(22).
A. Mesopleuron with precoxal groove deep and crenulate; Nearctic, rare.

B. Mesopleuron with precoxal groove not sculptured; Neotropical, rare.

Figure for couplet 23

24(22).
A. Forewing 3RSa less than 1.5 times as long as r (usually less than 1.2 times). Antenna usually with fewer than 25 flagellomeres.

B. Forewing 3RSa more than 1.6 times as long as r (usually more than 1.8 times). Antenna often with more than 25 flagellomeres.

Figure for couplet 24

25(16).
Propodeum coarsely sculptured. A. Forewing marginal cell short, with vein 3RSb reaching wing margin less than 0.8 times distance between apex of stigma and wing tip; forewing (RS+M)a straight; Neotropical, rare.

Propodeum smooth and shiny, at most with weak punctures at base of setae. B. Forewing marginal cell long, with vein 3RSb reaching wing margin more than 0.8 times distance between apex of stigma and wing tip; forewing (RS+M)a variable but often distinctly curved or angled posteriorly shortly after arising from 1M; widespread, common.

Figure for couplet 25

26(25).
A. Ventral border of clypeus sinuate in front view. Flagellum with fewer than 50 flagellomeres; southern USA and northern Mexico (one species, A. insolitus (Shenefelt) formerly the sole species of Alienoclypeus).

B. Ventral border of clypeus arched in front view. Flagellum usually with more than 60 flagellomeres; widespread, common.

Figure for couplet 26

27(26).
A. Mandibles massive; Neotropical, rare.

B. Mandibles of normal size; widespread, common.

Figure for couplet 27

28(27).
A. Metasoma with three very long pseudo-ovipositors (more than 5 times as long as body) extending from anal region (true ovipositor and ovipositor sheaths much shorter; arrows in images below); Neotropical, rare.

B. Metasoma without pseudo-ovipositors; widespread, common.

Figure for couplet 28

29(28).
A. Scape very long, more than 2.5 times as long as mid-width; Neotropical, uncommon.

B. Scape much shorter, less than 2.0 times as long as mid-width; widespread, common.

Figure for couplet 29

30(29).
A. Body long and slender. Metasomal terga 2-5 very long and slender, tergum 3 more than 2 times as long as maximum width: posterior margins of terga 3-5 strongly emarginate and membranous medially, with extremely long internal apodemes; Neotropical, rare.

B. Body not as above. Metasomal terga 2-5 never so long and slender, tergum 3 less than 2 times as long as maximum width; posterior margins of terga 3-5 at most weakly emarginate and never with a distinct membranous median zone, and with internal apodemes never more than 2 times as long as medially wide; widespread, common.

Figure for couplet 30

31(30).
A. Posterior halves of metasomal terga 3-5 thickly sclerotized and convex in profile. AA. Metasomal terga coarsely sculptured in most species.

B. Posterior halves of metasomal terga 3-5 less thickly sclerotized and straighter in profile. BB. Metasomal terga mostly smooth.

Figure for couplet 31

Generic treatments

Acgorium Sharkey & Quicke, 2020

Diagnosis. Propodeum smooth with a complete midlongitudinal carina; mesopleuron lacking a sculptured precoxal groove; tergum 1 with deep sculpture.

Biology. The single described species, A. felipechavarriai Sharkey, is a gregarious ectoparasitoid of Dyscophellus phraxanor Godman and Salvin (Hesperiidae) (Quicke et al. 2020b).

Diversity. Uncommon; one described species.

Distribution. The sole known species is from Costa Rica.

Publications. Quicke et al. (2020b).

Figure 1.
Figure 1. Acgorium felipechavarriai Sharkey.

Amyosoma Viereck, 1913

Diagnosis. Similar to Myosoma but hind femur and tibia with short setae. Median tergum 1 more than 2.0 times as long as maximum width. Scape shorter ventrally than dorsally (when antenna directed anteriorly).

Biology. Amyosoma chinense has been widely introduced to Neotropical areas to control Diatraea species (Crambidae) on sugarcane (Bennett 1971).

Diversity. Other than the introduced species, one species is described from the New World with probably more than 40 undescribed species.

Distribution. Previously thought to be restricted to the Old World but widespread in the Neotropics.

Publications. Papp (2012) described a new species from Honduras.

Figure 2
Figure 2. Amyosoma sp.

Atanycolus Foerster, 1862

Diagnosis. Raised median area of median tergite 1 not abruptly emarginate posterolaterally. Tergum 2 with triangular or (rarely) quadrate raised area narrowing posteriorly. Scape with false sub-apical margin, well-separated from the real margin, and scape basally abruptly and concavely narrowing.

Biology. Parasitoids of bark-boring beetles especially Cerambycidae and Buprestidae. Several species have been used to attempt the biological control of the emerald ash borer including Atanycolus cappaerti, a species described in Marsh et al. (2009). A. insolitus (Shenefelt 1978), formerly placed in the monotypic genus Alienoclypeus, is reported as a parasitoid of the weevil Scyphophorus Schoenherr.

Diversity. 65 described species worldwide; about 40 of these are in the New World most of which are Nearctic. We estimate undescribed species will more than double this number.

Distribution. Common in the Nearctic from Canada to Costa Rica; also common in the Palaearctic, but rarely in the Old World tropics (including Australia).

Publications. Shenefelt (1943) revised the North American species.

Note. Quicke et al. (2023) synonymized Alienoclypeus Shenefelt, 1978 with Atanycolus.

Figure 3
Figure 3. Atanycolus sp.

Bracon Fabricius, 1804

Diagnosis. Forewing vein 3RSa more than 1.6 times as long as crossvein r. Median tergite 1 not especially elongate, less than 2.0 times as long as greatest width. Marginal cell of fore wing long, vein 3RSb reaching wing margin at least 0.8 times distance between apex of stigma and wing tip. Scape shorter ventrally than dorsally.

Biology. Parasitoids of a wide range of Lepidoptera, Coleoptera, Diptera, and phytophagous Hymenoptera larvae. Flores et al. (2006) described the first phytophagous species. Many Neotropical species appear to be gregarious (Sharkey et al. 2021a).

Diversity. Approximately 200 species from the Neotropical and 90 from the Nearctic. About 1,000 species described worldwide, and many thousands more are undescribed. Very common and highly species-rich.

Distribution. Cosmopolitan.

Publications. Canadian and USA species were revised by Muesebeck (1925) as Microbracon; however, this is of limited use because of the large number of undescribed species. Sharkey et al. (2021a, b) described 48 species from Costa Rica and most of these were associated with host data.

Notes. Several Neotropical species groups are distinctive and some in particular are larger and/or more heavily sculptured than is typical of most Nearctic and Old World species; nevertheless, separation of these into distinct genera would almost certainly render the remainder of Bracon paraphyletic, which they probably are already.

Figure 4. Bracon sp.

Calobracon Szépligeti, 1902

Diagnosis. Antennal sockets protruding and distally acutely angulate in profile. Scape very short and distinctive, flared dorsally and ventrally.

Biology. Unknown.

Diversity. Two described species, fewer than 10 undescribed.

Distribution. Neotropical, to date known only from Argentina, Brazil, Paraguay and Peru.

Publications. Papp (2011) redescribed the type species.

Figure 5. Calobracon sp.

Cervellus Szépligeti, 1904

Diagnosis. First flagellomere with thorn-like, pre-apical projection.

Biology. Specimens of Cervellus piranga were reared from the papaya borer weevil Pseudopiazurus obesus associated with Carica papaya in Brazil (Penteado-Dias et al. 2007). Another species was reared from a Coleoptera larva (Quicke 1989c).

Diversity. Seven described species, perhaps as many undescribed.

Distribution. Tropical South America.

Publications. Fahringer (1930) provided a key to the species known at that time.

Figure 6. Cervellus sp.

Chartobracon van Achterberg, 1983

Diagnosis. Body very strongly dorsoventrally depressed, mesosoma more than 2.8 times as long as maximum height.

Biology. Chartobracon huggerti van Achterberg, has been reared from Callidium F. (Cerambycidae) tunnels under spruce bark.

Diversity. Two described species, only one in the Nearctic.

Distribution. Canada (1 species); also northern Europe.

Publications. Van Achterberg (1983a); Quicke and Sharkey (1989) described the Nearctic species.

Figure 7
Figure 7. Chartobracon sp.

Coeloides Wesmael, 1838

Diagnosis. Scape shorter ventrally than dorsally; flagellum uniformly dark, without white or cream-colored band at mid-length. Basal flagellomere expanded ventrally.

Biology. Parasitoids of bark-boring Coleoptera larvae, especially Scolytidae, Curculionidae and Buprestidae. Some species are probably important in the regulation of bark beetle pests of various coniferous trees.

Diversity. 32 described species, 13 New World species.

Distribution. Canada, USA, and Mexico; also Palaearctic.

Publications. Mason (1978) revised and keyed the species of USA and Canada.

Figure 8. Coeloides sp.

Compsobracon Ashmead, 1900

Diagnosis. Hypopygium truncated in profile. Most are large, brightly colored with long ovipositors and have thus been collected frequently.

Biology. One species has been reared from Crambidae (Quicke 1989c). Another was reared from an unidentified caterpillar boring in a thick branch of Alibertia concolor (Santos et al. 2004).

Diversity. 18 described species, perhaps twice or more that number undescribed.

Distribution. Southern USA (one record from Texas) to Brazil and Paraguay, Bahamas and Trinidad.

Publications. No comprehensive revision.

Figure 9. Compsobracon sp.

Compsobraconoides Quicke, 1989

Diagnosis. Propodeum simple, entirely smooth. Metasoma with median tergite 2 flat or weakly pinched anteromedially. Scape shorter ventrally than dorsally. Middle part of face clearly demarked from lateral parts by a pair of sub-medial, longitudinal carinae running ventrally from the antennal sockets. Metasomal tergite 2 without midbasal, posteriorly narrowing triangular area.

Biology. Yu and Quicke (1997) described a new species parasitizing adult Azteca ulei (Formicidae) in plant-formed domatia. The reared species described by Fortier & Nishida (2004) was transferred to Gozmanycomp (see Gozmanycomp treatment).

Diversity. At least 5 described species; a very variable genus as currently defined, some species have well-developed midlongitudinal grooves on median tergite 1, some have claws with rounded basal lobes, some have pale-striped antennae, some have the metasoma distinctly sinuous in dorsal view, and some have the suture between metasomal terga 2 and 3 strongly buckled medially. This is likely a polyphyletic group.

Distribution. Southern USA to South America and the Caribbean

Publications. No comprehensive treatment of the genus.

Figure 10. Compsobraconoides sp.

Cyanopterus Haliday, 1835

Diagnosis. Posterior halves of metasomal terga 3-5 mostly smooth, less thickly sclerotized, and relatively flat profile, compared to Digonogastra. Many species intermediate in these features  are difficult to place. Scape less than 2.0 times as long as mid-width. Scape apicolaterally emarginate and longer ventrally than dorsally.

Biology. Hosts include larvae of Coleoptera and Lepidoptera.

Diversity. Seven Nearctic and 25 Neotropical species described, many Nearctic and hundreds of Neotropical species undescribed.

Distribution. Nearctic and Neotropics. Common in the Nearctic and very common in the Neotropics. Also in Palearctic.

Publications. There is no comprehensive treatment of the genus in the New World.

Figure 11. Cyanopterus sp.

Cyclaulacidea Quicke and Delobel, 1995

Diagnosis. Face rugose with a lot of granulate sculpture, often with a raised granulate area medially. Raised median area of median tergite 1 rather square in cross-section, lateral margin more or less carinate posteriorly.

Biology. Parasitoids of Chrysomelidae (Bruchinae) and Curculionidae in palm fruit and similar situations (Quicke and Delobel 1995; Villemant and Simbolotti 2000).

Diversity. Eleven described species, many more undescribed.

Distribution. Neotropical: Mexico south to Brazil.

Publications. Quicke and Delobel (1995) erected the genus and included one species. Leathers et al. (2005) revised and keyed the species.

Figure 12. Cyclaulacidea sp.

Cyclaulax Cameron, 1911

Diagnosis. Length of tergum 2 about half the length of tergum 3. Scape longer ventrally than dorsally. Middle part of face clearly demarked from lateral parts by a pair of sub-medial, longitudinal carinae running ventrally from the antennal sockets. Metasomal tergum 2 without midbasal, posteriorly narrowing triangular area.

Biology. Unknown.

Diversity. Eleven described species, about 90 undescribed.

Distribution. Neotropical: Central America south to Brazil.

Publications. The nine Cyclaulax species represented by types and deposited in the Hungarian National Museum, Budapest, as well as the type-species of the genus, Cyclaulax grandiceps Cameron, housed in the Natural History Museum, London, were redescribed by Papp (2007a) who also presented an identification key to ten (of 11) species.

Figure 13. Cyclaulax sp.

Digonogastra Viereck, 1912

Diagnosis. Posterior halves of metasomal terga 3-5 partly sculptured, more thickly sclerotized, and relatively convex in profile, compared to Cyanopterus. Median tergite 1 usually with weak medial sub-posterior pit divided by midlongitudinal carina or with several sub-posterior, sub-medial pits. Many species intermediate in these features are difficult to place. Scape less than 2.0 times as long as mid-width, apicolaterally emarginate, longer ventrally than dorsally.

Biology. Hosts include larvae of Coleoptera and Lepidoptera, the latter includes pests of sugarcane and sorghum (Wharton et al. 1989, Quicke 1989a).

Diversity. 260 described species, perhaps several thousand undescribed species.

Distribution. New World, especially diverse in the Neotropics. Possibly extending into the eastern Palaearctic, and one reported from Africa,

Publications. Species descriptions are scattered in many publications and there is no comprehensive revision of the genus. Sharkey et al. (2021a) described five species from Costa Rica.

Notes. Many species were originally included in Iphiaulax Foerster, the latter is now redefined and restricted to the Old World (Quicke 1988c).

Figure 14. Digonogastra sp.

Gnathobracon Costa, 1864

Diagnosis. Mandibles massive.

Biology. Unknown.

Diversity. Two described species, probably a few others undescribed.

Distribution. Neotropical: French Guiana, Peru.

Publications. Quicke and Huddleston (1991) described the second species.

Figure 15. Gnathobracon sp.

Gozmanycomp Papp, 2007

Diagnosis. Propodeum smooth with a distinct longitudinal carina midposteriorly. Scape shorter ventrally than dorsally. Middle part of face clearly demarked from lateral parts by a pair of sub-medial, longitudinal carinae running ventrally from the antennal sockets. Metasomal tergum 2 without midbasal, posteriorly narrowing triangular area.

Biology. Fortier and Nishida (2004) described Gozmanycomp cinnamoni (as Compsobraconoides cinnamoni) reared from galls caused by larvae of the weevil, Camptocheirus sp. (Curculionidae), on branches of Cinnamonum cinnamomifolium (Lauraceae).

Diversity. Two described species, perhaps a dozen more undescribed.

Distribution. Neotropical: Costa Rica to Brazil.

Publications. Papp (2007b).

Figure 16. Gozmanycomp sp.

Gracilibracon Quicke, 1995

Diagnosis. Metasomal tergum 2 strongly 'pinched-up' anteromedially. Scape with false sub-apical margin, well-separated from the real margin, and scape basally abruptly and concavely narrowing

Biology. Unknown.

Diversity. Three described species.

Distribution. Neotropical.

Publications. Quicke (1995).

Figure 17. Gracilibracon sp.

Habrobracon Ashmead, 1895

Diagnosis. Forewing 3RSa less than 1.5 times length of r (usually less than 1.2 times). Antenna usually with fewer than 25 flagellomeres. Scape not, or only weakly, emarginate apicoIaterally. Scape shorter ventrally than dorsally. Forewing Crossvein r often slightly sinuate.

Biology. Hosts include concealed larvae of a wide variety of Coleoptera, Lepidoptera, Diptera and Hymenoptera but is best known for its gregarious parasitism of flour and grain moths. Habrobracon hebetor has been extensively studied in the laboratory.

Diversity. About 35 described species, and 14 in the New World. Perhaps two or three times these numbers yet to be described.

Distribution. Cosmopolitan.

Publications. Fischer (1968) provided a key to species. Kittel & Maeto (2018) restored H. brevicornis from synonymy.

Figure 18. Habrobracon sp.

Hemibracon Szépligeti, 1906

Diagnosis. Raised median area of tergum 1 semicircularly emarginate posterolaterally. Scape with false sub-apical margin, well-separated from the real margin, and scape basally abruptly and concavely narrowing.

Biology. Hemibracon rufidorsum was reared from Cobelura wappesi (Cerambycidae) (Murgas et al. 2018).

Diversity. 22 described species, perhaps several hundred more undescribed.

Distribution. Neotropical, Costa Rica to Argentina.

Publications. Papp (2011) provided a key to and redescribed the six species of Hemibracon described by Szépligeti.

Figure 19. Hemibracon sp.

Lapicida Quicke, 1989

Diagnosis. Propodeum and metanotum both with a complete lamelliform midlongitudinal carina. Precoxal groove deep and crenulate. Scape not, or only weakly, emarginate apicoIaterally and shorter ventrally than dorsally.

Biology. Associated with aquatic vegetation (Sagittaria).

Diversity. One described and one undescribed species are known. It is rarely collected because of its particular habitat.

Distribution. Canada to central USA

Publications. Quicke and Sharkey (1989).

Figure 20. Lapicida aquatica Quicke.

Lasiophorus Haliday, 1838

Diagnosis. Face with a median projection that is approximately round in cross section. Rather large specimens.

Biology. Unknown.

Diversity. Two described species and perhaps a dozen undescribed; older catalogues include more but these have subsequently been transferred to other genera.

Distribution. Neotropical: Mexico to Brazil.

Publications. Fahringer (1930) provided a key. Van Achterberg (1983b) re-diagnosed the genus, redescribed and illustrated the type species, and provided a key to the four species that were then included in the genus. Quicke (1989a) transferred two of these species to Palabracon.

Figure 21. Lasiophorus sp.

Leptobracon Szépligeti, 1901

Diagnosis. Metasomal terga 2-5 long and slender and tergum 3 more than 2 times as long as maximally wide: posterior margins of terga 3-5 strongly emarginate and membranous medially, with extremely long internal apodemes. A long, slender wasp.

Biology. Unknown.

Diversity. Four described species, few undescribed, rarely collected.

Distribution. Neotropical.

Publications. Fahringer (1930) provided a key to species.

Figure 22. Leptobracon sp.

Megabracon Szépligeti, 1906

Diagnosis. Scape very long, more than 2.5 times as long as mid-width. Scape apicolaterally emarginate, longer ventrally than dorsally, large specimens.

Biology. De Santis and Esquivel (1966) reported Megabracon as a parasitoid of Mallodon spinibarbis (as Stenodontes spinibarbis) (Cerambycidae).

Diversity. There are nine described species but none described since 1920. Probably 2 or 3 times this number of species are undescribed.

Distribution. Neotropical: Brazil to Guatemala.

Publications. There is no revision of the genus.

Figure 23. Megabracon sp.

Megacoeloides Quicke, 1989

Diagnosis. Scape longer ventrally than dorsally (when antenna directed anteriorly); flagellum with white or cream-colored band near midlength. Basal flagellomere expanded ventrally.

Biology. Unknown.

Diversity. One described species, probably fewer than 10 undescribed.

Distribution. Neotropical: Colombia, French Guiana, Peru.

Publications. Quicke (1989b).

Figure 24. Megacoeloides sp.

Myosoma Brullé, 1846

Diagnosis. Similar to Amyosoma but hind femur and tibia with long black setae. Metasomal tergum 1 more than 2.0 times as long as maximum width. Scape shorter ventrally than dorsally.

Biology. Parasitoids of Lepidoptera borers, especially Pyralidae (Mason 1978).

Diversity. 18 described species, probably 20 or 30 undescribed.

Distribution. Southern USA to Brazil.

Publications. Mason (1978) revised the species of North America.

Figure 25. Myosoma sp.

Myosomatoides Quicke, 1994

Diagnosis. Hind tibia strongly laterally compressed, very broad in profile with especially long setae. Metasomal tergum 2 with a midlongitudinal ridge.

Biology. One species has been reared from larvae of a stem-boring Diatraea sp. (Pyralidae) (Quicke 1994).

Diversity. Two described species and a few undescribed.

Distribution. Neotropical: widely distributed but rarely collected.

Publications. Quicke (1994).

Figure 26. Myosomatoides pennipes (Westwood).

Palabracon Quicke, 1988

Diagnosis. Flagellomere 1 simple or occasionally apically flared. Face with large, horizontal, dorsally concave plate-like projection.

Biology. One species has been reared from Diatraea (Lepidoptera: Crambidae) (Quicke 1988b).

Diversity. Three described species, probably a handful more undescribed, rarely collected.

Distribution. Neotropical: Brazil, Mexico, Ecuador.

Publications. Quicke (1988b).

Figure 27. Palabracon sp.

Pheloura van Achterberg, 1989

Diagnosis. Large specimens with three pseudo-ovipositors that are more than 5 times as long as body. The true ovipositor and ovipositor sheaths are much shorter.

Biology. Unknown.

Diversity. Only one species; rarely collected.

Distribution. Neotropical: Colombia, Costa Rica.

Publications. Van Achterberg (1989).

Figure 28. Pheloura sp.

Sacirema Quicke, 1995

Diagnosis. Scape shorter ventrally than dorsally. Middle part of face clearly demarked from lateral parts by a pair of sub-medial, longitudinal carinae running ventrally from the antennal sockets. Face coarsely carinate-rugose medially with little or no crenulate sculpture and without a raised granulate area medially. Raised median area of metasomal tergum 1 rounded in cross-section. Metasomal tergum 2 without midbasal, posteriorly narrowing triangular area, its length subequal to tergum 3 length.

Biology. Unknown.

Diversity. Three described species, dozens more undescribed.

Distribution. Neotropical.

Publications. Papp (2007c) described a new species from the Lesser Antilles, transferred a species from Bracon, redescribed the type species, and provided an illustrated key to the three known species. Sharkey et al. (2021b) described a new species from Costa Rica.

Figure 29. Sacirema sp.

Vipio Latreille, 1804

Diagnosis. Some species cannot be distinguished with certainty from Vipiomorpha; however, the key above works in more than 90% of the cases. Vipiomorpha always have an angular basal lobe on the claws whereas that is true only of approximately 50% of Vipio. Vipiomorpha never has a pair of long setae or small group of setae above the hypoclypeal depression, and never has an elongate labio-maxillary complex which is present in nearly all Vipio.

Biology. Parasitoids of concealed Lepidoptera, Coleoptera, and probably 'Symphyta'  (Hymenoptera) larvae.

Diversity. There are 11 described species in the Nearctic and 16 in the Neotropics with some species overlapping. There are probably 20 to 40 undescribed New World species.

Distribution. Canada to South America though less common in the southern portion of the range. Also Palaearctic, Palaeotropics (occasional),

Publications. Inayatullah et al. (1998) revised the Nearctic species north of Mexico. Quicke et al. (2020a) revised the species south of Nicaragua and included 13 species.

Figure 30. Vipio sp.

Vipiomorpha Tobias, 1962

Diagnosis. Propodeum coarsely sculptured. Forewing marginal cell short, 3RSb reaching wing margin less than 0.8 times distance between apex of stigma and wing tip, as in Vipio; forewing (RS+M)a straight. Scape not, or only weakly, emarginate apicoIaterally and shorter ventrally than dorsally

Biology. Unknown.

Diversity. Three described species, a few more undescribed.

Distribution. Only one undescribed species from South America known in the New World. Also Palearctic and Afrotropical (especially southern Africa).

Publications. Tobias (1962) erected the genus.

Figure 31. Vipiomorpha sp.

Acknowledgements

We thank Kees van Achterberg for permission to use and modify his drawings. Special thanks to our three referees: Alejandro Alejandro Zaldívar Riverón and Scott R. Shaw who thoroughly checked the key, and John Huber for very careful attention to the wording throughout. Finally, thanks to editors Heather Proctor and Spencer Monckton.

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