Michael Sharkey1*
Robert Wharton2
Kees van Achterberg3
1Hymenoptera Institute, 1339 La Loma Dr., Redlands, California, 92373, USA.
2Department of Entomology, Texas A&M University, College Station, Texas 77843, U.S.A. (formerly)
3Naturalis Biodiversity Center, Postbus 9517, 2300 RA Leiden, The Netherlands
* Corresponding author: [email protected]
Michael Sharkey1*
Robert Wharton2
Kees van Achterberg3
1Hymenoptera Institute, 1339 La Loma Dr., Redlands, California, 92373, USA.
2Department of Entomology, Texas A&M University, College Station, Texas 77843, U.S.A. (formerly)
3Naturalis Biodiversity Center, Postbus 9517, 2300 RA Leiden, The Netherlands
* Corresponding author: [email protected]
This is an updated overview of the New World (Western Hemisphere) members of Macrocentrinae. It is a revision of the keys by Wharton (1997) and Sharkey et al. (2021a) and contains a synopsis of each genus. The synopses include the following sections: diagnosis, biology, diversity, distribution, publications, and notes. There are four genera reported from the New World.
Wharton, R.A. 1997. Subfamily Macrocentrinae, pp.310-313. In: Wharton, R.A., Marsh, P.M., and Sharkey, M.J. (eds). Manual of the New World Genera of the Family 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
Macrocentrinae is a moderately diverse subfamily containing four genera in the New World. Most of the described species belong to the genus Macrocentrus Curtis. Ahlstrom (2005) revised the species of North America north of Mexico, updating Muesebeck (1932). The latest generic synopsis is by van Achterberg (1993). Shaw and Huddleston (1991) provide a useful summary of the literature, with special reference to biology. New World specimens of Braconidae can be identified to subfamily using the key by Sharkey et al. (2023). Readers using that key will be automatically directed here if they obtain an identification of Macrocentrinae. The present work includes a key to the four New World genera, and a synopsis of each genus. The synopses include the following sections: diagnosis, biology, diversity, distribution, publications, and notes. Images are by Sharkey. General morphological terminology can be found in Sharkey et al. (2023) and in the Hymenoptera Anatomy Ontology Portal (http://portal.hymao.org/projects/32/public/ontology/).
Phylogeny
No phylogenetic analyses at the genus level have been conducted. In a molecular phylogenetic study, Sharanowski et al. (2011) recovered Macrocentrinae as sister to Xiphozelinae and these were found to be sister to the clade Amicrocentrinae + Charmontinae.
Biology
The Macrocentrinae include both solitary and gregarious koinobiont endoparasitoids of lepidopterous larvae (Clausen, 1940; Shaw and Huddleston, 1991). Where known, the gregarious species are polyembryonic. Detailed biological information is available for two species, M. ancylivorus Rohwer (Daniel, 1932; Finney et al., 1947) and M. cingulum Brischke (as referred to in recent literature van Achterberg, 1993; Farahani et al., 2012) or with authorship incorrectly attributed to Reinhard or as M. grandii Goidanich (Parker, 1931; Ding et al., 1989) or as M. gifuensis Ashmead (a misidentification) in some of the more important earlier publications. Although only a single individual of M. ancylivorus emerges from its host, initial development is polyembryonic (Daniel, 1932). Details of the development of other macrocentrines are largely unknown. It is possible, however, that those species that emerge as solitary parasitoids (at least in Macrocentrus and Hymenochaonia Dalla Torre) are initially polyembryonic in their development. New World macrocentrines have been reared from 10 families of Lepidoptera (Yu et al, 2016; Sharkey et al. 2021a).
Many of the macrocentrines are pale colored and crepuscular to nocturnal. The species of Macrocentrus and Hymenochaonia have long ovipositors, often laying their eggs in early instars of hosts concealed in leaf whorls, stems, tubers, or rolled leaves. Austrozele Roman and Dolichozele Viereck, on the other hand, have short ovipositors, and the available records indicate that they attack exposed caterpillars, e.g., Arctiidae and Noctuidae.
Common genera
Macrocentrus and Hymenochaonia are the major New World genera. There are about 190 described species of Macrocentrus worldwide, and about one-fourth of these occur in the New World. The remaining 46 species of described macrocentrines are divided amongst seven other genera worldwide. We estimate that there are several hundred or more undescribed New World species.
Distribution
Cosmopolitan.
Distinguishing features
Macrocentrines can be recognized by the absence of an occipital carina and the presence of small teeth on the hind trochantellus (Fig. 1). With the exception of one known species, M. incompletus Muesebeck, all described species have three submarginal cells.
Note: In the generic treatments we give rough estimates of the number of undescribed species that may occur in the New World, e.g., “There are 26 BINs (proxies for species) from Costa Rica on the Barcode of Life Data System (BOLD, Nov. 25, 2024), and we estimate that there may be a hundred or more undescribed species in the New World.” To arrive at these very conservative numbers we used estimates of the number of trees in Costa Rica compared to the remainder of the New World. Cazzolla Gatti et al. (2022) estimated there to be 40,123 species of trees in the New World. Nelson Zamora (personal communication), the leading Costa Rican botanist, estimates there to be 2,500 species of trees in Costa Rica, or ~6.2% New World fauna. Over the last ten years Dan Janzen, Winnie Hallwachs and their team have barcoded Malaise trap samples from all over Costa Rica, making it the best sampled country on the Barcode of Live Data System database, though it is far from complete. In the example above, if we were to assume that the 26 BINs represented 6.2% of the New World fauna we would arrive at an estimate of 419 species for the New World. However, we refrain from this detailed estimate and instead make the generalized and conservative estimate of “a hundred or more.”
Table 1. List of New World genera of Macrocentrinae
Diagnosis. Petiole with laterope deep (couplet 2B); ovipositor short, about equal to apical height of metasoma (Fig. 2A).
Biology. Solitary parasitoids of Noctuidae (van Achterberg, 1993) and Geometridae (Sharkey et al., 2021a).
Diversity. There are 22 described species worldwide, two in the Nearctic and four in the Neotropical regions. Presently (Nov. 25, 2024) there are six BINs (proxies for species) from Costa Rica and 12 BINs from Canada on the Barcode of Life Data System (BOLD). There may be close to 100 undescribed species in the New World.
Distribution. Cosmopolitan.
Publications. Ahlstrom (2005) revised the Nearctic species occurring north of Mexico. Sharkey et al. (2021a) described two new species from Costa Rica.
Diagnosis. Ovipositor much shorter than metasoma (Fig. 3A). Laterope weakly impressed or absent (couplet 1A).
Biology. Solitary parasitoids of Erebidae (Arctiidae in the literature) and Noctuidae (Sharkey et al., 2021a).
Diversity. Nine described species worldwide, with three in the Nearctic and three in the Neotropical regions. There may be several dozen more undescribed in the Neotropics.
Distribution. Nearctic, Neotropical, western Canada (British Columbia) south to Brazil.
Publications. Ahlstrom (2005) revised the Nearctic species occurring north of Mexico. Sharkey et al. (2021a) described two new species from Costa Rica.
Diagnosis. Ovipositor as long as or longer than the metasoma, usually at least as long as the body (Fig. 4A). Laterope weakly impressed or absent (couplet 1A).
Biology. Reared primarily from Pyralidae and Tortricidae. Allen (1962) summarized the biology of H. delicata (Cresson) (as Macrocentrus delicatus). Sharkey et al. (2021a) described species reared from Erebidae (Arctiidae in the literature), Depressariidae, Pyralidae, Crambidae, and Gelechiidae.
Diversity. 23 described species worldwide. There are 26 BINs (proxies for species) from Costa Rica on the Barcode of Life Data System (BOLD, Nov. 25, 2024), and we estimate that there may be a hundred or more undescribed species in the New World.
Distribution. Holarctic, Neotropical (southern Canada to Argentina). Introduced into Europe for biological control of the Oriental fruit moth, Grapholita molesta (Busck), but not recovered.
Publications. Ahlstrom (2005) revised the Nearctic species occurring north of Mexico. Sharkey et al. (2021a) described 13 new species from Costa Rica
Diagnosis. Ovipositor as long as or longer than the metasoma, usually at least as long as the body (Fig. 5A). Laterope deeply impressed (couplet 1B). Most of the small specimens (less than 5 mm.) belong to this genus.
Biology. Most of the detailed work has been on polyembryonic species (overview in Shaw and Huddleston, 1991). Tortricidae and Pyralidae are the most frequently recorded hosts, however Sharkey et al. (2021a, b) recorded 16 species with Crambidae as hosts and also two with Depressariidae as hosts.
Diversity. About 210 species described worldwide, with 46 in the Nearctic and 29 in the Neotropical regions. There are 25 BINs (proxies for species) from Costa Rica and 35 BINs from Canada on the Barcode of Life Data System (BOLD, Nov. 25, 2024), and we estimate that there are several hundred undescribed species in the New World.
Distribution. Cosmopolitan.
Publications. Ahlstrom (2005) revised the Nearctic species occurring north of Mexico. Sharkey et al. (2021a, b) described 21 new species from Costa Rica.
We thank A.P. Ranjith and Caroline Boudreault for their reviews of the submitted manuscript and the editors of CJAI for their efforts.
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Allen, H.W. 1962. Parasites of the Oriental fruit moth in the Eastern United States. United States Department of Agriculture Technical Bulletin, 1265: 25-26.
Clausen, C.P. 1940. Entomophagous Insects. McGraw-Hill, New York and London. 688pp.
Daniel, D.M. 1932. Macrocentrus ancylivorus Rohwer, a polyembryonic braconid parasite of the Oriental fruit moth. New York State Agricultural Experiment Station Technical Bulletin, 187: 1-101.
Ding, D., Swedenborg, P.D., and Jones, R.L. 1989. Chemical stimuli in host-seeking behavior of Macrocentrus grandii (Hymenoptera: Braconidae). Annals of the Entomological Society of America, 82: 232-236. https://doi.org/10.1093/aesa/82.2.232
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Finney, G.L., Flanders, S.E., and Smith, H.S. 1947. Mass culture of Macrocentrus ancylivorus and its host, the potato tuber moth. Hilgardia, 17: 437-483.
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Parker, H.L. 1931. Macrocentrus gifuensis Ashmead, a polyembryonic braconid parasite in the European corn borer. United States Department of Agriculture Technical Bulletin, 230: 1-62. https://doi.org/10.22004/ag.econ.163054
Sharanowski, B.J., Dowling, A.P., and Sharkey, M.J. 2011. Molecular phylogenetics of Braconidae (Hymenoptera: Ichneumonoidea), based on multiple nuclear genes, and implications for classification. Systematic Entomology, 36: 549–572. https://doi.org/10.1111/j.1365-3113.2011.00580.x
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
Sharkey, M.J., Baker, A., McCluskey, K., Smith, A., Naik, S., Ratnasingham, S., Manjunath, R., Perez, K., Sones, J., D’Souza, M., Jacques, B.S., Hebert, P., Hallwachs, W., and Janzen, D. 2021b. Addendum to a minimalist revision of Costa Rican Braconidae: 28 new species and 23 host records. ZooKeys, 1075: 77–136. https://doi.org/10.3897/zookeys.1075.72197
Sharkey, M., Athey, K.J., Fernández-Triana, J.L., Penteado-Dias, A.M., Monckton, S.K., and Quicke, D.L., 2023. Key to the New World subfamilies of the family Braconidae (Hymenoptera: Ichneumonoidea). Canadian Journal of Arthropod Identification, 49: 1-43. https://doi.org/10.3752/cjai.2023.49
Shaw, M.R and Huddleston, T. 1991. Classification and biology of braconid wasps (Hymenoptera: Braconidae). Handbooks for the Identification of British Insects, 7: l-126. Available online: https://www.royensoc.co.uk/wp-content/uploads/2022/01/Vol07_Part11.pdf
van Achterberg, C. 1993. Revision of the subfamily Macrocentrinae Foerster (Hymenoptera: Braconidae) from the Palaearctic region. Zoologische Verhandelingen Leiden, 286: 1-110. Available online: https://repository.naturalis.nl/pub/317629/ZV1993286001.pdf
Wharton, R.A. 1997. Subfamily Macrocentrinae, pp.310-313. In: Wharton, R.A., Marsh, P.M., and Sharkey, M.J. (eds). Manual of the New World Genera of the Family Braconidae (Hymenoptera). Special Publication of the International Society of Hymenopterists, 439 pp.