Aerophilus Szépligeti, 1902

Diagnosis. The structure of the attachment of the metasoma to the mesosoma (Fig. 1) is unique among New World agathidines. The metasomal foramen is widely separated from the coxal cavities and there is a transverse ridge between them. The structure is shared only by Braunsia Kriechbaumer, an Old World genus.

Biology. Members attack a wide range of lepidopteran families, e.g., Batrachedridae, Coleophoridae, Crambidae, Elachistidae, Gelechiidae, Pyralidae, Thyrididae, and Tortricidae.

Diversity. There are an estimated 300 species globally, of which only about 60 are described.

Distribution. Widespread in the New World in both tropical and temperate regions. Worldwide in distribution, but far less diverse in the Old World.

Publications. Sharkey et al. (2016) revised the eastern North American species and included all USA and Canadian species in a key. Sharkey et al. (2011a) revised many Costa Rican species of this genus under the name Lytopylus, a name that was later found to be erroneously applied.

Note. Most described species were formerly included in Bassus.

Agathirsia Westwood, 1882

Diagnosis. Very similar to some species of Crassomicrodus. All Crassomicrodus have simple claws without a basal lobe, whereas most Agathirsia have claws with a basal lobe (Fig. 4B). The ovipositors of Crassomicrodus specimens are short, barely exserted from the apex of the metasoma (couplet 32A); those of Agathirsia are at least as long as half the length of the metasoma (couplet 32B). The mouthparts (galea) of Agathirsia specimens are often elongate (as in couplet 14A), whereas those of Crassomicrodus are not elongate (as in couplet 14B).

Biology. Bibby (1961) cites the larva of Acontia cretata (Grote & Robinson) (Noctuidae) as a host of A. testacea Muesebeck. This is the only host record for the genus. The fact that this host pupates in the soil and that all members of Agathirsia have thick mandibles suited for digging prompted Pucci and Sharkey (2004) to suggest that all species attack members of Noctuidae that pupate in the soil.

Diversity. Thirty-two species are described and there may be an additional 30 species still undescribed.

Distribution. Only known from Mexico and southwestern USA although some species are likely to occur in dry areas of other countries of Central America.

Publications. Pucci and Sharkey (2004) revised the species.

Note. The monophyly of Agathirsia and Crassomicrodus is uncertain.

Agathis Latreille, 1804

Diagnosis. Frons not bordered by carina posteriorly (couplet 11b). Head usually elongate and mouthparts (galea) elongate (couplet 14A and Fig. 5A). Hind coxal cavities open to metasomal foramen, or narrowly closed (couplet 15A). Tarsal claws with or rarely without a basal lobe (couplet 4B).

Biology. Lepidopteran host families include the following: Gelechiidae, Coleophoridae, Oecophoridae, Tortricidae, Heliodinidae, Pyraustidae, Cochylidae, Momphidae, Epermeniidae and Incurvariidae, particularly those feeding in flower heads (Simbolotti and van Achterberg, 1999).

Diversity. This is a hard number to determine; 46 species from the West Palearctic are considered valid (Simbolotti and van Achterberg, 1999) but molecular methods were not used to delineate species. There are about seven described Nearctic species from the USA and Canada, but many more undescribed. There are a number of Neotropical species placed in Agathis, but none of these belongs to the genus, e.g., A. alvarengai de Santis and A. amoena Brullé.

Distribution. Agathis is primarily Holarctic and Oriental with a few undescribed species found in Costa Rica, Colombia, and Chile. In the Nearctic they are most diverse in the southwestern USA.

Publications. Muesebeck’s (1927) key is the only one available and it conflates about seven genera treated under Agathis. It also treats only a small fraction of the species. In essence, there is no practical way to identify New World species of Agathis.

Alabagrus Enderlein, 1920

Diagnosis. Frons bordered by carinae posteriorly (couplet 11A). First metasomal tergum usually convex or with a median carina (rarely with a pair of carinae that converge medially, but then with other irregular longitudinal striae (couplet 13BB and 13BBB). First metasomal tergum mildly to acutely convex (Fig. 7).

Biology. Exclusively parasitoids of Crambidae.

Diversity. 192 described species and an estimated 800 undescribed species.

Distribution. Restricted to the New World and primarily Neotropical. Alabagrus occurs from southern Canada to Argentina.

Publications. Sharkey (1988) revised the genus and included about 100 species. With the advent of sequence data, it was discovered that many species definitions in this revision were actually species complexes. Leathers and Sharkey (2003) revised the species of La Selva, Costa Rica, and this too is replete with incorrectly circumscribed species. Sharkey et al. (2018) revised the species of Costa Rica concentrating on reared and barcoded specimens from Guanacaste province and those species are accurately delineated.

Note: Astiria Enderlein, 1920, Craspedobothrus Enderlein, 1920, and Liyptia Enderlein, 1920 are junior synonyms of Alabagrus.

Amputoearinus Sharkey, 2006

Diagnosis. Gena acute posteroventrally (couplet 29A). RS+M vein of forewing complete and tubular (Fig. 8C), as in Earinus. Propleuron with a distinct protuberance (couplet 29A). First metasomal tergum smooth and flat (Fig. 8E). Notauli absent (Fig. 8D).

Biology. Amputoearinus niger Lindsay & Sharkey was reared from an undescribed species of Dysodia, Thyrididae (Sharkey et al. 2011a).

Diversity. Sixteen described species and approximately 30 undescribed species.

Distribution. Restricted to the Neotropics and recorded from Mexico to Peru and Brazil.

Publications. Lindsay and Sharkey (2006) revised the genus, which was erected by Sharkey (2006).

Aphelagathis Sharkey, 2015

Diagnosis. First median tergite smooth lacking microsculpture and carinae (Fig. 9F). Tarsal claws simple with a large basal lobe (couplet 4B). Notauli deeply impressed and lacking sculpture (Fig. 9B). Frons not margined by carinae (Couplet 11B).

Biology. Hosts are known for two species, and both are parasitoids of grass-feeding Hesperiidae.

Diversity. Eleven described species, an estimated 10 undescribed species.

Distribution. Widespread in the southern half of the USA, south to Costa Rica and the Caribbean islands. Aphelagathis appears to be most diverse in the southwestern USA and northwestern Mexico.

Publications. Sharkey et al. (2015) revised the species.

Note. The sole species described before 2015 was formerly included in Bassus.

Chilearinus Sharkey, 2022

Diagnosis. Notauli absent (Fig. 10C). Second submarginal cell quadrate (Fig. 10B). Cub vein of hind wing present (Fig. 10D). Subbasal cell of hind wing angled where Cub vein intersects (Fig. 10). Tarsal claws with basal lobe (couplet 4B).

Biology. Unknown

Diversity. Hundreds of species of which only five are described.

Distribution. Chile and southern Argentina with a few species in high altitudes as far north as Colombia and Ecuador.

Publications. Sharkey et al. (2022) erected the genus, described two new species, and included other species formerly placed in Earinus.

Crassomicrodus Ashmead, 1900

Diagnosis. Very similar to some species of Agathirsia. All species of Crassomicrodus have simple claws without a basal lobe (Fig. 12A), whereas most Agathirsia have simple claws with a basal lobe (Fig. 12B). The ovipositors of Crassomicrodus specimens are short, barely exerted from the apex of the metasoma (Fig. 11A); those of Agathirsia are at least as long as half the length of the metasoma (couplet 32B). The mouthparts (galea) of Agathirsia specimens are often elongate (as in couplet 14A), whereas those of Crassomicrodus are not elongate (as in couplet 14B).

Biology. Autographa californica (Speyer) (Noctuidae) attacked by C. fulvescens (Cresson) is the unique host record for the genus.

Diversity. There are 15 described species and about the same number may be undescribed.

Distribution. Primarily North and Central America, but extending to southern Canada, with one record from northwestern Colombia. Most diverse in Mexico and southwestern USA.

Publications. Figueroa et al. (2011) revised the genus.

Note. The monophyly of Agathirsia and Crassomicrodus is uncertain.

Cremnops Foerster, 1863

Diagnosis. Malar length at least half the eye height (Fig. 13A). Frons with lateral carinae (Fig. 13B). Notaulus impressed (Fig. 13C). Second submarginal cell quadrate (Fig. 13D). Tarsal claws cleft with basal pectination (Fig. 13E). Propodeum and hind coxa lacking granulate sculpture (couplet 8B).

Biology. Members of the family Crambidae are the usual hosts; other recorded hosts include Tortricidae and Sesiidae, but they may be erroneous.

Diversity. Seventy-seven species are described, 33 of which are found in the New World. There may be another 30 undescribed species worldwide.

Distribution. Worldwide with the highest diversity in temperate North America.

Publications. Tucker et al. (2015) revised the New World species.

Note. Žikić et al. (2011) proposed the genus Neocremnops for a group of New World species with asymmetrical tarsal claws, a character elucidated in Tucker et al. (2015). I do not recognize the genus because there is no evidence that the remaining species of Cremnops form a monophyletic group.

Earinus Wesmael, 1837

Diagnosis. RS+M vein of forewing complete or almost so (Fig. 14B). Gena rounded posteroventrally. The genus does not occur in the Neotropics.

Biology. Reared from the following families: Coleophoridae, Geometridae, Gracillariidae, Noctuidae, and Tortricidae.

Diversity. Eighteen described species, only five in the New World, and perhaps 20 undescribed species.

Distribution. Primarily temperate Holarctic with several species in the Oriental realm. Restricted to Canada, the USA and northern Mexico (Sonora) in the New World. As the genus name suggests, specimens of the genus are exclusively found in the spring.

Publications. Sharkey et al. (2022) revised the Nearctic species and added two new species. Braet (2002) described two new species of that he assigned to Earinus, however they both belong to Lytopylus and were transferred by Sharkey et al. (2022). Berta (2000) described a new species and presented a key to three Neotropical species. These were transferred to Chilearinus and Lytopylus by Sharkey et al. (2022).

Note. Previously, Sharkey et al. (2006) included a diverse clade from south temperate South America in the genus. However, Sharkey et al. (2022) erected Chilearinus to refer to this group.

Gelastagathis Sharkey, 2015

Diagnosis. Clypeus flared anteriorly and mouth appearing cyclostome-like (Fig. 15D).Temples bulging behind eyes (Fig. 15E).

Biology. Unknown, however the strong rounded mandibles are reminiscent of Agathirsia and Crassomicrodus, therefore soil dwelling caterpillars are the suspected hosts.

Diversity. Two described species and probably fewer than 10 undescribed.

Distribution. Known from the two specimens, one from Texas and one from Massachusetts.

Publications. Sharkey and Chapman (2015) described the genus and the two included species.

Hemichoma Enderlein, 1920

Diagnosis. Claws cleft (couplet 4A). Ovipositor shorter than metasoma (Fig. 16A). Mesoscutum smooth, lacking notauli (Fig. 16B). Gena expanded posteroventrally (couplet 6A).

Biology. The few known hosts are in the family Erebidae.

Diversity. Seven described species, 10-20 more undescribed.

Distribution. Neotropics.

Publications. Meierotto et al. (2019b) described several species from Costa Rica and included the first host records.

Labagathis Enderlein, 1920

Diagnosis. Tarsal claws cleft (couplet 4A). Ovipositor longer than metasoma (Fig. 17A). Propodeum with granulate sculpture (couplet 8A). Gena elongate (Fig. 17A).

Biology. Unknown

Diversity. L. rufoatra Enderlein is the sole described species and it may be the only species.

Distribution. Neotropical: Known from Colombia, Venezuela, Peru, and Suriname

Lytopylus Foerster, 1862

Diagnosis. Mesoscutum smooth, lacking notauli (Fig. 18D). Claws simple with a basal lobe (couplet 4B). Third tergum smooth (Fig. 18D). Second submarginal cell of forewing large, lacking a petiole (Fig. 18E). Gena rounded posterolaterally (couplet 21B).

Biology. Recorded as parasitoids of the following families: Gelechiidae, Depressariidae, Oecophoridae.

Diversity. 39 described species and several hundred undescribed species.

Distribution. Restricted to the New World, from the northeastern USA south to Argentina, primarily Neotropical.

Publications. Kang et al. (2017) revised reared species from Costa Rica. Braet (2002) treated a number of Neotropical species, which he assigned to Earinus, however most of these belong to Lytopylus and were transferred by Sharkey et al. (2022).

Note. The revision by Sharkey et al. (2011a) employed the name Lytopylus in error and later Sharkey et al. (2016) transferred all of species erroneously described under Lytopylus in that paper to Aerophilus Szépligeti, 1902. Over past decades, Lytopylus was considered a junior synonym of Bassus Fabricius, Microdus Nees, or Agathis Latreille. Sharkey et al. (2016) removed the genus from synonymy and synonymized Agathellina Enderlein, 1920, Ditropia Enderlein, 1920, and Austroearinus Sharkey, 2006 under Lytopylus.

Marjoriella Sharkey, 1983

Diagnosis. Tarsal claws simple with a basal lobe (couplet 4B). Ovipositor shorter than metasoma (Fig. 19A). 2RS2 vein of forewing long and strong (Fig. 19D).

Biology. Unknown

Diversity. Four described species and probably a handful more undescribed.

Distribution. Rainforests of tropical South America.

Publications. Sharkey (1983) revised the species.

Note. Characters unique to members of Marjoriella are the projections that emanate ventrally from the propleura and mesopleura (Fig. 19C).

Mesocoelus Schulz, 1911

Diagnosis. Forewing venation greatly reduced (Fig. 20B). Hind legs elongate (Fig. 20A).

Biology. The sole recorded host is Acrocercops sp., Gracillariidae (Muesebeck, 1932).

Diversity. Three described species, about 10 undescribed.

Distribution. Recorded only from Caribbean islands and Brazil in the literature (Achterberg, 1990) but widespread in the Neotropics, e.g., Costa Rica, Colombia.

Publications. Bortoni and Penteado-Dias (2015) described a new species and presented a key to the three described species.

Neothlipsis Sharkey, Parys and Clutts, 2011

Diagnosis. First tergum partly or completely granulate and lacking prominent pair of longitudinal carinae (couplet 22A). Third tergum lacking longitudinal striae, usually partly granulate (Fig. 21B). Notauli pitted (Fig. 21B). Gena not elongate (Fig. 21A). Frons not bordered by carinae or grooves posteriorly (couplet 11B).

Biology. Recorded as parasitoids of the following families: Gelechiidae, Crambidae, Tortricidae, Elachistidae, Coleophoridae.

Diversity. Ten described species and perhaps another 40 undescribed.

Distribution. Canada south to Costa Rica and the Lesser Antilles.

Publications. Sharkey et al. (2011b) erected the genus, transferred nine species from other genera and described a new species. Sharkey et al. (2021) described a species from Costa Rica.

Note. Members of this genus were formerly included in Bassus.

Pharpa Sharkey, 1986

Diagnosis. Tarsal claws simple with a basal lobe (couplet 4B). Frons bordered posteriorly with carinae (couplet 11A); hind coxa lacking granulate sculpture (couplet 12B). First tergum smooth with two widely spaced converging carinae forming a tear-shaped basal area (couplet 13A, Fig. 22D).

Biology. Unknown, possibly Crambidae as in its sister group Alabagrus.

Diversity. Three described species and about a dozen undescribed.

Distribution. Neotropical, southern Mexico to northern Argentina.

Publications. Sharkey (1986) erected the genus and included three species.

Plesiocoelus van Achterberg, 1990

Diagnosis. Forewing venation reduced (Fig. 23B), but not as extensively as that of Mesocoelus (couplet 1A). Metasomal tergum 1 with granulate or strio-granulate sculpture (Fig. 23C).

Biology. Depressariidae (rearing record in Janzen’s Costa Rican database (http://janzen.sas.upenn.edu/caterpillars/database.lasso)

Diversity. Four described species however not all of them belong in this genus. Bortoni and Penteado-Dias (2015) described two new species under Plesiocoelus however they both belong to other genera. Plesiocoelus anomalus Bortoni & Penteado-Días belongs in Therophilus [Therophilus anomalus (Bortoni & Penteado-Días, 2015) n. comb.] The evidence for this is in their figure 11, which shows the elongate Cub vein emanating from an angle in the basal cell of the hind wing. Plesiocoelus areolatus Bortoni & Penteado-Días belongs in Aerophilus, [Aerophilus areolatus (Bortoni & Penteado-Días, 2015) n. comb.] The evidence for this is in their figure 19, which shows the elevated medial portion of the first metasomal median tergite. There are probably about 10 undescribed species in the Neotropics.

Distribution. Rare but widespread in the Neotropics. Plesiocoelus also occurs in Southeast Asia (see below).

Publications. Achterberg (1990) erected the genus and included one species. Long and Achterberg (2016) described a new species from Vietnam. Bortoni and Penteado-Dias (2015) reported a range extension for P. bassiformis van Achterberg.

Pneumagathis Sharkey 2015

Diagnosis. Claws simple with a basal lobe (couplet 4B). Frons not bordered by carinae or grooves posteriorly (couplet 11B). Propodeal spiracle elongate (Fig. 24D).

Biology. Parasitic on a wide variety of grass feeding Hesperiidae (Janzen et al., 1998 [as Bassus]).

Diversity. Two described species, and probably fewer than 5 species undescribed, if any.

Distribution. Widespread in the Nearctic, Mexico, and Central America.

Publications. Sharkey and Chapman (2015) erected the genus and included two species. Janzen et al. (1998) elucidated the biology and host range of Pneumagathis erythrogastra Cameron (as Bassus brooksi Sharkey).

Sesioctonus Viereck, 1912

Diagnosis. Claws simple lacking a basal lobe (couplet 4A). Mesoscutum smooth (Fig. 25B). Notauli lacking (couplet 30A, Fig. 25B).

Biology. The single host record is from Erebidae (Viereck, 1912b)

Diversity. 32 described species and about 70 undescribed.

Distribution. Widespread but rarely collected in the Neotropics from Mexico to Brazil.

Publications. Briceño (2003) revised the genus. Sharkey and Briceño (2005) added five new species. Sulca and Sharkey (2012) added three new species and updated the key.

Smithagathis Sharkey, 2017

Diagnosis. Notauli absent (Fig. 26C). Hind coxal cavity open (couplet 26A). Claws with basal lobe (couplet 4B). R1 of forewing curved and reaching margin far from apex of wing (Fig. 26F). Second submarginal cell small and petiolate (Fig. 26F).

Biology. Unknown.

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

Distribution. Known only from the type locality, which has an altitude of 3,660 meters in the Colombian Andes.

Publications. Sharkey and Chapman (2017) erected the genus and described the sole species.

Therophilus Wesmael, 1837

Diagnosis. Posterior apex of scutellum with a distinct depression in the form of a semicircle or two distinct pits (couplet 23A). First tergum not granulate, rarely striogranulate (couplet 22B). Cub vein of hind wing long and partly tubular. Apical margin of subbasal cell angled (couplet 19A). Notauli impressed and sculptured (Fig. 27D). Claws with a basal lobe (couplet 4B);

Biology. Parasites of Tortricidae and Elachistidae.

Diversity. 44 described species, probably 200 undescribed species.

Distribution. Worldwide, in the New World the genus is more species rich in the Nearctic region.

Publications. Sharkey and Stoelb (2012) resurrected the genus, described 11 new species and transferred 33 species from Bassus, Agathis and Microdus.

Note. Most species were formerly included in Bassus.

Trachagathis Viereck, 1913

Diagnosis. Frons bordered by grooves posteriorly (couplet 11A). Hind coxa with granulate sculpture (Fig. 28B). Second cubital sell of forewing small (Fig. 28C). Notauli impressed and sculptured (couplet 20A). Claws simple with a basal lobe (couplet 4B). Second submarginal cell reduced to a small point, rarely absent (Fig. 28C).

Biology. The sole host record is on Pyralidae.

Diversity. Three described species and about 10 undescribed species.

Distribution. Widespread in the Neotropical region.

Publications. Sharkey (2007) revised the genus.

Zacremnops Sharkey and Wharton, 1985

Diagnosis. Claws cleft (couplet 4A). Forewing areolet quadrate, not or only slightly narrower anteriorly (Fig. 29B). Ovipositor as long as or longer than half the length of the metasoma (Fig. 29A). Body predominantly black (Fig. 29A). Frons not bordered by a carina posteriorly (couplets 11B).

Biology. Four species are known to be parasitoids of Crambidae.

Diversity. Five described species and perhaps about five undescribed.

Distribution. Southern USA to Argentina.

Publications. Sharkey (1990) revised the genus, synonymized several species, and presented a key. Sharkey et al. (2021) presented host data and illustrations for four species.

Zamicrodus Viereck, 1912

Diagnosis. Propleuron with a distinct protuberance; gena expanded into an acute angle posteroventrally (couplet 21A). Notauli smooth, unsculptured (Fig. 30B). First tergum completely smooth and lacking pair of longitudinal carinae (Fig. 30B). Frons not bordered by carinae or grooves posteriorly (couplet 11B). Claws simple, with distinct basal lobe (couplet 4B).

Biology. There are two records of Z. brianbrowni Sharkey on Quasimellana sethos (Mabille) (Hesperiidae).

Diversity. Three described species, perhaps about eight undescribed species.

Distribution. Widespread from Costa Rica south to Brazil.

Publications. Viereck (1912a) and Brullé (1846) each described a species. Sharkey et al. (2021) described a new species and included the first host records.

Zelomorpha Ashmead, 1900

Diagnosis. Gena not modified into a flange posteriorly (couplet 4B). Forewing areolet triangular or if quadrate much narrower anteriorly (Fig. 31B). Ovipositor shorter than half the length of metasoma (Fig. 31A, C). Claws cleft (couplet 4A).

Biology. The following families are recorded as hosts of Zelomorpha species: Erebidae, Euteliidae, Lasiocampidae, Noctuidae, Nolidae, Notodontidae.

Diversity. 74 species described and hundreds more are undescribed. Some Old World species are assigned to Zelomorpha, but these all belong other genera such as Disophrys Förster, Biroia Szépligeti, and Coccygidium Saussure.

Distribution. Southern USA to Brazil.

Publications. Descriptions are scattered in numerous publications. Meierotto et al. (2019a) revised 15 species from Costa Rica, included host records, and transferred all described species placed in other genera to Zelomorpha (Meierotto et al. 2019b). Sarmiento et al. (2004) described the first known gregarious species of Zelomorpha (as Coccygidium).