PROC. ENTOMOL. SOC. WASH.
124(1), 2022, pp. 75–84
A NEW SPECIES OF OPIUS (BELLOPIUS) (HYMENOPTERA: 
BRACONIDAE) REARED FROM HEXACHAETA (DIPTERA: 
TEPHRITIDAE) FLIES IN COLOMBIA, WITH A KEY TO NEW WORLD 
SPECIES OF OPIUS PARASITIC ON FRUIT-INFESTING TEPHRITIDAE
Robert R. Kula, Helmuth Aguirre, and Pedro Alexander Rodriguez-Clavijo
(RRK) Systematic Entomology Laboratory, Beltsville Agricultural Research 
Center, Agricultural Research Service, U.S. Department of Agriculture, c/o 
National Museum of Natural History, Smithsonian Institution, P.O. Box 37012, 
MRC-168, Washington, DC 20013-7012, USA (e-mail: Robert.Kula@usda.gov, 
urn:lsid:zoobank.org:author:76A42D9B-5EFF-4494-8729-C9DF9915CA5B); 
(HA, PARC) Instituto Colombiano Agropecuario (ICA), C.I. Tibaitatá km 14 vía 
Bogotá-Mosquera, Colombia (e-mail: helmuthaguirre@gmail.com, urn:lsid:zoobank.
org:author:F1C9B312-15D7-4A7D-A544-0FEDC6C5515D)
Abstract.—A new species of fruit fly parasitoid, Opius (Bellopius) hexachaeta 
Kula and Aguirre (Hymenoptera: Braconidae), is described. The parasitoid was 
reared from larvae of the genus Hexachaeta (Diptera: Tephritidae) feeding on seeds in 
entireleaf spiritweed, Aegiphila integrifolia (Jacq.) B.D. Jacks (Lamiales: Lamiaceae). 
The morphology and taxonomic placement of the new species is discussed.
Key Words: Opiinae, taxonomy
DOI: 10.4289/0013-8797.124.1.75
Bellopius Wharton (Braconidae: sulcus absent or nearly so (Wharton and 
Opiinae) is a subgenus of Opius Wesmael Marsh 1978), wings heavily infumate, 
that consists of 11 species (Yu et al. 2016) forewing 2cu-a reduced, and pronope 
known only from the Neotropical Region present. Wharton (1997) proposed the 
(Wharton and Yoder 2021). One species, subgenus Bellopius within Opius for spe-
Opius (Bellopius) bellus Gahan, has been cies of the bellus group.
reported from Thailand (Chinajariyawong As far as is known, species of Bellopius 
et al. 2000); however, this is the only are koinobiont endoparasitoids of fruit flies 
report of a Bellopius species outside the (Diptera: Tephritidae) (Wharton 1997), 
Neotropical Region and thus requires although host use is known only for two 
confirmation (Wharton and Yoder 2021). species: O. (B.) bellus and Opius (B.) hirtus 
The species currently in Bellopius were Fischer. Host records for both species have 
placed in Desmiostoma Förster in Fischer been complied in Yu et al. (2016) and col-
(1977) and included in a key to the New lectively span six genera; the vast majority 
World species of that genus. Wharton of hosts are species of Anastrepha Schiner. 
(1983) recognized a group of species in Notably, O. (B.) bellus has been reported 
Desmiostoma (i.e., bellus group) with the from at least 22 host species (all but three 
following combination of features and of them Anastrepha spp.) collectively from 
transferred them to Opius: notauli and at least 27 host plants, including pests such 
mesonotal midpit absent and precoxal as Anastrepha fraterculus (Wiedemann), 
Downloaded From: https://bioone.org/journals/Proceedings-of-the-Entomological-Society-of-Washington on 17 Oct 2022
Terms of Use: https://bioone.org/terms-of-use	Access provided by Smithsonian Institution
76 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Anastrepha obliqua (Macquart), Anastrepha out in locations of Cundinamarca depart-
serpentina (Wiedemann), Anastrepha ment. The host plant was photographed 
striata Schiner, and Ceratitis capitata in situ using a Samsung A50 cell phone. 
(Wiedemann). However, O. (B.) bellus may Subsequently, fruits were placed in plas-
be a species complex considering its host tic containers (15 cm height, 15 cm diam-
range, patterns of host use, geographic dis- eter) under laboratory conditions in a 
tribution, and variation with respect to color Memmert IN55 Incubator with 24 hours 
(Ovruski et al. 2000). Further, Wharton dark, 14–19°C temperature, and 60–75% 
(1997) noted at least two undescribed spe- relative humidity. Adult flies were fed 
cies of Bellopius, but additional species have once a day for 2–3 days with a mixture of 
not been described or assigned to Bellopius water, sugar, and hydrolyzed protein. Fly 
since that time. Species richness and pat- larvae fed upon the same fruit on which 
terns of host use are unclear for Bellopius they were collected. A host fly puparium 
and will involve disentangling species com- was dissected and yielded an opiine bra-
plexes, as well describing new species and conid pupa (Fig. 3). All opiines emerged 
documenting new hosts. from host puparia within 12–47 days of 
The Instituto Colombiano Agropec- being placed in laboratory conditions. 
uario (ICA) is currently monitoring the Nine female and seven male parasitoids 
geographic distribution of tephritid fruit emerged and were placed in vials with 
flies in Colombia via the Programa 96% alcohol. Six female and three male 
Nacional Moscas de la Fruta (ICA-PNMF), specimens were placed in a bath of 99% 
which employs the following three sam- ethanol for 24 hours and then a bath of 
pling techniques: (1) McPhail traps baited formaldehyde for five minutes; those 
with hydrolyzed protein for collecting specimens were then air dried for 10 min-
Anastrepha spp., (2) Jackson traps baited utes, point-mounted, and used to describe 
with trimedlure for attracting C. capitata the new species. The remaining three 
and also with both methyl eugenol and cue- females and four males were not used to 
lure for attracting species of Bactrocera write the description due to their poor 
Macquart and Dacus Fabricius, and (3) physical condition and are not part of the 
sampling of exotic and native fruit for estab- type series. We have retained those spec-
lishing plant-insect associations. Larvae of imens in 96% alcohol as genetic vouch-
Hexachaeta Loew were found, using the ers that are deposited in Museo Javeriano 
latter approach, feeding on seeds in fruit de Historia Natural “Lorezo Uribe, S.J.”, 
of Aegiphila integrifolia (Jacq.) B.D. Jacks. Bogota, Colombia (MPUJ).
(Lamiales: Lamiaceae), entireleaf spirit- Host plant identifications were made 
weed (Figs. 1, 2). The larvae were para- by Ms. Guadalupe Caicedo at the her-
sitized by braconids (Fig. 3), which were barium of the Jardín Botánico de Bogotá 
identified as a new species of Bellopius. We José Celestino Mutis (JBB). Insect speci-
describe the new species herein and pro- mens were examined using a Nikon SMZ-
vide a key for identifying the New World 1000 stereomicroscope. The specimens 
species of Opius reported as parasitoids of of Braconidae were identified as Opius 
tephritids that infest fruit (Yu et al. 2016). (Bellopius) using Wharton (1997); they 
were determined as a new species using 
diagnostic information and illustrations in 
Materials and Methods Fischer (1977), Wharton and Marsh (1978), 
Hexachaeta larvae were collected and Wharton and Yoder (2021), as well as 
using fruit sampling techniques carried through comparison with specimens of 
Downloaded From: https://bioone.org/journals/Proceedings-of-the-Entomological-Society-of-Washington on 17 Oct 2022
Terms of Use: https://bioone.org/terms-of-use	Access provided by Smithsonian Institution
VOLUME 124, NUMBER 1 77
Figs. 1–3. Opius (Bellopius) hexachaeta ex Hexachaeta sp. in fruit of Aegiphila integrifolia. 1, Fruit on host 
plant, A. integrifolia. 2, Hexachaeta sp. larva feeding on A. integrifolia seed. 3, Opius (B.) hexachaeta pupa and 
Hexachaeta sp. puparium.
described species. The first author (RRK) system. The following abbreviations are 
evaluated multiple images of all described used in the descriptions: head length (HL), 
species of Bellopius, as well as Opius head width (HW), temple width (TW), 
(Nosopaeopius) downesi Gahan, taken of face width (FW), face height (FH), eye 
specimens identified by braconid special- length (EL), eye height (EH), flagellomere 
ist Dr. Robert A. Wharton (retired, Texas 1 length (F1L), flagellomere 2 length 
A&M University). Holotypes were exam- (F2L), mesosoma length (ML), mesoscu-
ined for O. (B.) bellus, Opius (B.) chro- tum width (MW), mesosoma height (MH), 
maticus Fischer, Opius (Nosopaeopius) scutellar sulcus length (SSL), scutellar sul-
baldufi Muesebeck, and O. (N.) downesi. cus width (SSW), tergum 1 length (T1L), 
All type specimens of the new species are tergum 1 width (T1W), and terga one 
deposited in MPUJ. through eight (t1…t8).
Measurements were taken as described Terminology for anatomical features, 
in Wharton (1977) with additions and surface sculpture, and setation follows 
modifications as in Kula and Zolnerowich Sharkey and Wharton (1997). The mater-
(2005, 2008) using the software incorpo- ial examined section is formatted as in 
rated in a monitor Nikon DS-L3 camera Kula and Zolnerowich (2008). The key 
Downloaded From: https://bioone.org/journals/Proceedings-of-the-Entomological-Society-of-Washington on 17 Oct 2022
Terms of Use: https://bioone.org/terms-of-use	Access provided by Smithsonian Institution
78 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
to species of Opius known to parasitize carinae are absent in Opius (B.) cuzco-
fruit-infesting Tephritidae is based, in part, ensis Fischer. The gena, mesonotum, 
on the key in Wharton and Marsh (1978). propodeum, and metapleuron are setif-
Images of the new species and its host erous, but not notably dense (i.e., setose) 
were taken with a Nikon DS-L3 camera in O. (B.) hexachaeta (Figs. 4, 5, 8, 11); 
connected to a Nikon SMZ-1000 stere- they are setose in O. (B.) cuencaensis and 
omicroscope. Images were edited using Opius (B.) johannis Fischer.
Adobe Photoshop with the goal of ren- Description.—Female (Fig. 4). Body 
dering backgrounds of the unedited pho- length: 3.4–3.9 mm. Head (Figs. 5–7): HL 
tographic images more uniform in color 0.5–0.6X HW, HW 4.5–4.7X TW, FW 1.7–
and tone. For Fig. 11 two photos were 1.8X FH, EL 0.5–0.6X EH, F1L 1.3X F2L; 
combined to generate one high quality antenna with 40 flagellomeres, maxillary 
figure, and some setae were painted back palpus with 6 palpomeres, labial palpus 
in for clarity. The layout of individual fig- with 4 palpomeres; face smooth, setif-
ures into plates was accomplished using erous; frons smooth, glabrous; gena and 
Adobe Illustrator. vertex smooth, setiferous; occiput smooth, 
glabrous; eye glabrous; clypeus flat with 
Results and Discussion ventral margin lobed mesally, setiferous, 
labrum concealed by clypeus when man-
Opius (Bellopius) hexachaeta Kula and dible closed; mandible 2-toothed, smooth 
Aguirre, new species and setiferous, setae about as long as 
http://zoobank.org/3607871A-0ED0-4835- width of mandible at its midpoint, upper 
8C27-6B3B36433CC7 tooth longer than lower tooth.
Mesosoma (Figs. 5, 8, 11): ML 1.2–
(Figs. 4–11) 1.3X MW, ML 0.9–1.0X MH, MW 
Diagnosis.—Forewing vein 3RSb not 0.7–0.8X MH, SSL 0.2X SSW; pronotal 
complete to wing margin as tubular vein collar smooth and shiny, setiferous, setae 
in O. (B.) hexachaeta (Fig. 9); it is always oriented anteriorly; pronope large, lat-
complete to wing margin as tubular vein eral portion of pronotum smooth, mostly 
in all other species of Opius (Bellopius). glabrous but setiferous along margins, 
The mesosoma is entirely dark brown setae oriented anteriorly, anterolateral 
(nearly black) in O. (B.) hexachaeta (Figs. furrow shallow but present in dorsal 
4, 5, 8); it is at least partially or entirely one-fourth and crenulate from pronotal 
orange in Opius (B.) barrosensis Fischer, collar; notauli absent; mesoscutal midpit 
Opius (B.) bellus Gahan (Figs. 15, 16), absent; mesoscutum smooth and shiny, 
Opius (B.) campinaensis Fischer, Opius uniformly setiferous except two glabrous 
(B.) chromaticus Fischer, Opius (B.) cin- areas posterolaterally; scutellar sulcus 
gulaticornis Fischer, Opius (B.) fiebrigi crenulate, with three to four longitudinal 
Fischer, Opius (B.) hirtus Fischer (Fig. 17), carinae; scutellar disc smooth, setiferous, 
and Opius (B.) marcapatanus Fischer. All setae oriented posteriorly; propodeum 
legs (excluding trochantelli and bases of smooth except complete median carina, 
femorae) are entirely dark brown (nearly setiferous; epicnemial carina absent; 
black) in O. (B.) hexachaeta (Fig. 4); they precoxal sulcus present, smooth; poste-
are entirely yellowish orange with tarsi rior mesopleural furrow entirely smooth; 
brown in Opius (B.) cuencaensis Fischer. mesopleuron smooth, setiferous except 
The propodeum has a median carina in glabrous areas dorsad precoxal sulcus and 
O. (B.) hexachaeta (Fig. 11); propodeal posterodorsally dorsad episternal scrobe; 
Downloaded From: https://bioone.org/journals/Proceedings-of-the-Entomological-Society-of-Washington on 17 Oct 2022
Terms of Use: https://bioone.org/terms-of-use	Access provided by Smithsonian Institution
VOLUME 124, NUMBER 1 79
Figs. 4–10. Opius (Bellopius) hexachaeta, female. 4, Lateral habitus. 5, Head and mesosoma, lateral. 6, Head, 
anterior. 7, Head, dorsal. 8, Mesonotum, dorsal. 9, Forewing (arrow = 3RSb incomplete). 10, Hind wing.
metapleuron smooth, setiferous, setae ori- middle of stigma; vein length ratios: 2RS 
ented posteroventrally. 0.6–0.7X 3RSa, 2RS 0.5X 2M, 3RSa 0.7X 
Forewing (Fig. 9): Length 4.0–4.3 mm; 2M; veins C+Sc+R, 1RS, 1M, (RS+M)
stigma ellipitcal, distal margin tapering a, r, 2RS, 3RSa, 2M, 1m-cu, 1CUa, 1CUb, 
into vein R1; vein r arising slightly basad 2CUa, 1cu-a, 1-1A, and 2-1A complete 
Downloaded From: https://bioone.org/journals/Proceedings-of-the-Entomological-Society-of-Washington on 17 Oct 2022
Terms of Use: https://bioone.org/terms-of-use	Access provided by Smithsonian Institution
80 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 11. Opius (Bellopius) hexachaeta, female, propodeum and metasoma in dorsal view (arrows: a = median 
propodeal carina, b = smooth tergum 2).
Downloaded From: https://bioone.org/journals/Proceedings-of-the-Entomological-Society-of-Washington on 17 Oct 2022
Terms of Use: https://bioone.org/terms-of-use	Access provided by Smithsonian Institution
VOLUME 124, NUMBER 1 81
Figs. 12–15. Opius (Bellopius) spp. 12, 13, Opius (B.) downesi. 12, Head, anterior (arrow = basal tooth present). 
13, Metasoma, dorsal (arrow = striate tergum 2). 14, 15, Opius (B.) bellus. 14, Head, anterior (arrow = basal tooth 
absent). 15, Head and mesosoma, dorsal.
and tubular; veins (RS+M)b and r-m incomplete, nebulous, 1st subdiscal cell 
complete but not tubular; vein 3RSb not open.
complete to wing margin as tubular vein, Hindwing (Fig. 10): Veins C+Sc+R, 
proximal portion tubular ~1/3 distance Sc+R, R, R1, r-m, 1M, M+CU, cu-a, and 
to apical wing margin with remainder 1A complete and tubular, basal and sub-
spectral to wing margin; vein 3M incom- basal cells closed; RS minutely tubular 
plete, tubular proximally transitioning proximally and 2M tubular proximally 
to spectral distally to wing margin; vein along half its length, transitioning distally 
2CUb incomplete, nebulous; vein 2cu-a to nebulous and then spectral.
Downloaded From: https://bioone.org/journals/Proceedings-of-the-Entomological-Society-of-Washington on 17 Oct 2022
Terms of Use: https://bioone.org/terms-of-use	Access provided by Smithsonian Institution
82 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 16. Opius (Bellopius) bellus, female, lateral habitus.
Fig. 17. Opius (Bellopius) hirtus, head and mesosoma in dorsal view.
Metasoma (Fig. 11): T1L 0.8–0.9X posterolaterally; t3–t7 smooth, setiferous 
T1W, T2L 0.2X T2W, T3L 0.2–0.3X posteriorly with setae in single transverse 
T3W, T4L 0.3X T4W, T5L 0.1–0.4X T5W, line; t8 smooth, setiferous.
T6L 0.2–0.4X T6W, T7L 0.1X T7W; Color: Head (including antenna) entirely 
ovipositor longer than gaster; t1 smooth dark brown (nearly black), mandible dark 
with dorsal carinae extending slightly brown with yellowish orange tips, palpi 
posteriad spiracles, dorsope present, set- yellow; mesosoma entirely dark brown 
iferous laterally and posteriorly but oth- (nearly black); wings infumate, stigma 
erwise glabrous; t2 smooth, setiferous and veins brown; legs dark brown (nearly 
Downloaded From: https://bioone.org/journals/Proceedings-of-the-Entomological-Society-of-Washington on 17 Oct 2022
Terms of Use: https://bioone.org/terms-of-use	Access provided by Smithsonian Institution
VOLUME 124, NUMBER 1 83
black) except trochantellus and narrow than the gaster in O. (B.) cuencaensis and 
band at base of femora yellowish orange; O. (B.) cuzcoensis. Also, setation of the 
metasoma entirely orangish yellow. gena, mesonotum, propodeum, and meta-
Male: As in female except: Body pleuron is less dense in O. (B.) hexachaeta 
length: 2.5–3.0 mm. Head: HW 4.8–4.9X compared with the other three species, 
TW, FW 1.6–1.8X FH, EL 0.6–0.7X although the difference is less profound in 
EH; antenna with 38–42 flagellomeres. O. (B.) cuzcoensis. Additionally, while the 
Mesosoma: ML 1.1X MH; propodeum legs of both O. (B.) hexachaeta and O. (B.) 
entirely dark brown (almost black) to johannis are entirely dark brown except the 
mostly dark brown but with small testa- yellowish orange trochantelli and band at 
ceous area dorsomesally. base of each femora, the femorae of O. (B.) 
Host.—Opius (B.) hexachaeta was johannis are yellow apically where they 
reared from two new species of Hexachaeta articulate with the tibiae.
(Rodríguez et al. in prep.) with larvae feed- Opius (B.) hexachaeta is the only spe-
ing on seeds in fruit of Aegiphila integri- cies of Bellopius with 3RSb not complete 
folia. Opius (B.) hexachaeta is the only to the wing margin as a tubular vein. Vein 
species of Braconidae reported from a 3RSb extends to the wing margin but is 
species of Hexachaeta. spectral for most of its length.
Material examined.—Holotype female: 
COLOMBIA: Cundinamarca: Anolaima, Key to the New World Species  
Vereda Santo Domingo, near to finca of Opius Reared from Fruit- 
Villa Mariana 4.80171°N, 74.47542°W, Infesting Tephritids
1532 m, fruits collected 1 Jan 2018, adults 
and parasitoids emerged 12–16 Feb P. A. 1.  Mandible with basal tooth (Fig. 12); t2 
Rodriguez; El Colegio, Predio Boyacá, distinctly striate (Fig. 13) ................... 2
4.55000°N, 74.48751°W, 861 m, fruits col-  Mandible with ventral surface evenly 
lected 3 Jan 2018, adults and parasitoids curved, basal tooth absent (Fig. 14); t2 
emerged 15–23 Jan 2019, P. A. Rodriguez smooth (Fig. 11) ................................. 3
(Museo Javeriano de Historia Natural 2(1).  Ovipositor shorter than gaster, t1 usually 
“Lorezo Uribe, S.J.”, MPUJ). Paratypes: rugulose or coriaceous ................ Opius 
Two females and one male, same data as (Nosopaeopius) baldufi ....... Muesebeck
holotype. Three females and two males,  Ovipositor at least as long as gaster, t1 
El Colegio, Predio Boyacá, 4.55000°N, usually striate (Fig. 13) ................Opius 
74.48751°W, 861 m, fruits collected 3  ........... (Nosopaeopius) downesi Gahan
Jan 2019, adults and parasitoids emerged 3(1).  Forewing vein 3RSb not complete to 
15–23 Jan 2019, P. A. Rodriguez (MPUJ). wing margin as tubular vein (Fig. 9); 
Etymology.—The stem of the specific mesosoma entirely dark brown (Figs. 4, 
epithet refers to the genus name of the 5, 8) ....... Opius (Bellopius) hexachaeta 
host fruit fly, Hexachaeta.  ............. Kula and Aguirre, new species
Discussion.—Opius (B.) hexachaeta  Forewing vein 3RSb complete to wing 
is most similar morphologically to O. (B.) margin as tubular vein; mesosoma at 
cuencaensis, O. (B.) cuzcoensis, and O. least partially yellow (Fig. 15) ............ 4
(B.) johannis; these four species have the 4(3)  Head, excluding ocellar triangle, yellow 
mesosoma entirely dark brown. In addition (Fig. 15); legs yellow except metatibia 
to the features listed above in the diagno- and metatarsus brown in some speci-
sis, the ovipositor is longer than the gaster mens (Fig. 16) ............................. Opius 
in O. (B.) hexachaeta, while it is shorter  .......................(Bellopius) bellus Gahan
Downloaded From: https://bioone.org/journals/Proceedings-of-the-Entomological-Society-of-Washington on 17 Oct 2022
Terms of Use: https://bioone.org/terms-of-use	Access provided by Smithsonian Institution
84 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
 Head entirely dark brown (Fig. 17); legs distribution records. Proceedings of the Ento-
(excluding trochantelli and bases of mological Society of Washington 110: 1–60.
femorae) entirely brown (cf. Fig. 4) ...... Ovruski, S., M. Aluja, J. Sivinski, and R. Whar-
ton. 2000. Hymenopteran parasitoids on 
 ...........Opius (Bellopius) hirtus Fischer fruit-infesting Tephritidae (Diptera) in Lat-
in America and the southern United States: 
Acknowledgments diversity, distribution, taxonomic status and 
their use in fruit fly biological control. Inte-
We thank the personnel of the Instituto grated Pest Management Reviews 5: 81–107.
Colombiano Agropecuario, Programa Sharkey, M. J. and R. A. Wharton. 1997. Morphol-
Nacional Moscas de la Fruta (ICA-PNMF). ogy and terminology, pp. 19–37. In Wharton, 
R. A., P. M. Marsh, and M. J. Sharkey, eds. 
We are grateful to Taina Litwak (USDA- Manual of the New World Genera of the 
ARS-SEL) for processing images and con- Family Braconidae (Hymenoptera). Special 
structing plates. Mention of trade names or Publication No. 1. International Society of 
commercial products in this publication is Hymenopterists, Washington, DC. 439 pp.
solely for the purpose of providing specific Wharton, R. A. 1977. New World Aphaereta spe-
cies (Hymenoptera: Braconidae: Alysiinae), 
information and does not imply recom- with a discussion of terminology used in the 
mendation or endorsement by the USDA. tribe Alysiini. Annals of the Entomological 
USDA is an equal opportunity provider and Society of America 70: 782–803. https://doi.
employer. org/10.1093/aesa/70.5.782.
Wharton, R. A. 1983. Variation in Opius hirtus 
Fischer and discussion of Desmiostoma Foer-
Literature Cited ster (Hymenoptera, Braconidae). Proceedings 
of the Entomological Society of Washington 
Chinajariyawong, A., A. R. Clarke, M. Jirasurat, 85: 327–330.
S. Kritsaneepiboon, H. A. Lahey, S. Vijay- Wharton, R. A. 1997. Generic relationships of 
segaran, and G. H. Waiter. 2000. Survey opiine Braconidae (Hymenoptera) parasitic 
of Opiine parasitoids of fruit flies (Diptera: on fruit-infesting Tephritidae (Diptera). Con-
Tephritidae) in Thailand and Malaysia. The tributions of the American Entomological 
Raffles Bulletin of Zoology 48: 71–101. Institute 30: 1–53.
Fischer, M. 1977. Hymenoptera: Braconidae Wharton, R. A. and P. M. Marsh. 1978. New 
(Opiinae II—Amerika). Das Tierreich 97: World Opiinae (Hymenoptera: Braconidae) 
1–1001. parasitic on Tephritidae (Diptera). Journal of 
Kula, R. R. and G. Zolnerowich. 2005. A new the Washington Academy of Sciences 68: 
species of Epimicta Förster (Hymenoptera: 147–167.
Braconidae) from North America and new Wharton, R. A. and M. J. Yoder. 2021. Parasitoids 
distribution records for Epimicta griffithsi of fruit-infesting Tephritidae. (http://paroffit.
Wharton. Proceedings of the Entomological org). [Last accessed 2 February 2021.].
Society of Washington 107: 78–83. Yu, D. S. K., C. van Achterberg, and K. Horstmann. 
Kula, R. R. and G. Zolnerowich. 2008. Revi- 2016. Taxapad 2016 – World Ichneumonoidea 
sion of New World Chaenusa Haliday sensu 2015. Taxonomy, Biology,  Morphology and 
lato (Hymenoptera: Braconidae: Alysiinae), Distribution. USB Flash drive. www.taxapad.
with new species, synonymies, hosts, and com, Nepean, Ontario, Canada.
Downloaded From: https://bioone.org/journals/Proceedings-of-the-Entomological-Society-of-Washington on 17 Oct 2022
Terms of Use: https://bioone.org/terms-of-use	Access provided by Smithsonian Institution