Genera of New World subfamilies of Braconidae (Hymenoptera): single-genus subfamilies Acampsohelconinae, Apozyginae, Dirrhopinae, Masoninae, and Meteorideinae
Michael Sharkey1*
,
Diana Carolina Arias Penna2
,
Cornelis van Achterberg3
,
James B. Whitfield4
,
Donald L. J. Quicke5
1Hymenoptera Institute, 1100 Pepperhill Circle, Lexington, Kentucky 40502, USA
2Bogotá, D.C., 111221, Cundinamarca, Colombia
3Naturalis Biodiversity Center, Postbus 9517, 2300 RA Leiden, Netherlands
4Department of Entomology, University of Illinois, Champaign, Illinois, USA
5Department of Biology, Faculty of Life Sciences, Chulalongkorn University, Bangkok, Thailand
* Corresponding author: [email protected]
Genera of New World subfamilies of Braconidae (Hymenoptera): single-genus subfamilies Acampsohelconinae, Apozyginae, Dirrhopinae, Masoninae, and Meteorideinae
Genera of New World subfamilies of Braconidae (Hymenoptera): single-genus subfamilies Acampsohelconinae, Apozyginae, Dirrhopinae, Masoninae, and Meteorideinae
Michael Sharkey1*
,
Diana Carolina Arias Penna2
,
Cornelis van Achterberg3
,
James B. Whitfield4
,
Donald L. J. Quicke5
1Hymenoptera Institute, 1100 Pepperhill Circle, Lexington, Kentucky 40502, USA
2Bogotá, D.C., 111221, Cundinamarca, Colombia
3Naturalis Biodiversity Center, Postbus 9517, 2300 RA Leiden, Netherlands
4Department of Entomology, University of Illinois, Champaign, Illinois, USA
5Department of Biology, Faculty of Life Sciences, Chulalongkorn University, Bangkok, Thailand
* Corresponding author: [email protected]
Abstract
This is an overview of the five New World (Western Hemisphere) braconid subfamilies that include only a single genus: Acampsohelconinae (Urosigalphus Ashmead), Apozyginae (Apozyx Mason), Dirrhopinae (Dirrhope Foester), Masoninae (Masona van Achterberg), and Meteorideinae (Meteoridea Ashmead). Synopses for each genus are updates or revisions from the treatments in Wharton et al. (1997) and include sections on phylogeny, biology, diversity, distribution, diagnosis, and important publications.
Wharton, R.A., Marsh, P.M., and Sharkey, J.J. (eds.) 1997. Manual of the New World Genera of Braconidae (Hymenoptera). Special Publication of the International Society of Hymenopterists. 439 pp.
Introduction
This publication is part of a series that updates the Manual of New World Braconidae (Wharton et al. 1997) and includes the five braconid subfamilies that have only a single genus represented in the New World. The key by Sharkey et al. (2023) may be used to confirm a subfamily identification. Readers using that key will be automatically directed here if they obtain an identification of Acampsohelconinae, Apozyginae, Dirrhopinae, Masoninae, or Meteorideinae.
Urosigalphus Ashmead is the only member of Acampsohelconinae found in the New World. It was included in Helconinae in the original version of the Manual (Sharkey, 1997b). Acampsohelconinae was erected by Tobias (1987) for a fossil (Acampsohelcon Tobias) in Baltic amber. Van Achterberg (2002) added three extant genera to the subfamily, i.e., Afrocampsis van Achterberg & Quicke, Canalicephalus Gibson, and Urosigalphus. Gibson's (1972a) key is useful for the identification of species in the United States and Canada; however, his keys to the Central American (Gibson 1972b, 1982a), and South American (Gibson 1974, 1982b) fauna treat only a small fraction of the species. Arias-Penna and Whitfield (2012) give a complete introduction to all aspects of the subfamily and revised the Colombian species.
The updated overview of Apozyginae in the New World is a revision of the treatment by Sharkey (1997a). Apozyginae contains one species, Apozyx penyai Mason. Mason (1978) proposed a new family, Apozygidae, for this species and later described the previously unknown female of the same species (Mason 1987).
The updated overview of Dirrhopinae in the New World is a revision of the treatment by Whitfield (1997a). Van Achterberg (1984) erected a new subfamily for Dirrhope which had been placed in several different subfamilies previously. Only a single relatively rare genus, Dirrhope Foester, 1851 is known.
The updated overview of Masoninae in the New World is a revision of the treatment by van Achterberg (1997). Two tribes were recognized by van Achterberg (1995): Mannokeraiini (restricted to Australia) and Masonini (widespread). Mannokeraiini has since been shown to belong to the Euphorinae (Belshaw and Quicke, 2002; Sharanowski et al., 2011; Stigenberg et al., 2015).
The updated overview of Meteorideinae in the New World is a revision of the treatment by Sharkey (1997c). Meteorideinae is a small subfamily with two genera, Meteoridea Ashmead and Pronkia van Achterberg. Only Meteoridea is found in the New World. Van Achterberg (1990) proposed the genus Pronkia and presented a key to separate members of Pronkia from those of Meteoridea.
General morphological terminology can be found in Sharkey et al. (2023). More detailed information is in the morphology chapter of the New World manual (Sharkey and Wharton, 1997) and in the Hymenoptera Anatomy Ontology Portal (http://portal.hymao.org/projects/32/public/ontology/). All images are by M. Sharkey unless noted otherwise.
Taxonomic treatments
ACAMPSOHELCONINAE
M. Sharkey and D.C. Arias-Penna
Urosigalphus Ashmead, 1889
Phylogeny. Urosigalphus (Fig. 1) is currently placed in Acampsohelconinae, although it was traditionally included in the tribe Brachistini of Helconinae. A recent molecular phylogeny based on ribosomal DNA (28S and 18S) and nuclear protein-coding genes (CAD and ACC) elevated Brachistini to subfamily level and suggested that Urosigalphus (and presumably other acampsohelconines, which were not included in the analyses) is sister to a large clade including all helconoids (Helconinae, Brachistinae) and the macrocentroid subcomplex. This subcomplex is within the helconoid complex and includes Orgilinae, Homolobinae, Microtypinae, Charmontinae, Amicrocentrinae, Xiphozelinae, and Macrocentrinae (Sharanowski et al. 2011). This placement was also recovered by Jasso-Martínez et al. (2022).
Biology. Unknown for all other genera but members of Urosigalphus are endoparasitoids of the larval stage, perhaps egg-larval parasitoids, of Curculionidae and Bruchinae (Chrysomelidae). A complete list of the host records for Urosigalphus is contained in Table 1 of Arias-Penna and Whitfield (2012).
Diversity. There are 104 described species (Yu et al., 2016; Arias-Penna and Whitfield, 2012), but many more are undescribed.
Distribution. Urosigalphus is restricted to and widespread in the Nearctic and Neotropical realms. Afrocampsis (Afrotropical) and Canalicephalus (Indo-Australian) are Old World genera.
Diagnosis. (Fig. 2). The hind claws are dimorphic, i.e., the outer claw is longer than the inner claw. There are sometimes other differences between the two claws as well, e.g., different curvature and the presence of a basal lobe on the inner claw. Vein Cub of the hind wing is present. The metasoma is in the form of an unsegmented carapace. The second submarginal cell and crossvein 2cu-a of the forewing are absent.
Publications. Gibson (1972a, b, 1974, 1982a, b) described and keyed the New World species of Urosigalphus. Arias-Penna (2007) reported on the altitudinal range and geographic distribution of the genus in Colombia as well as the variation in body length and coloration among New World species. Arias-Penna and Whitfield (2012) described 23 new species from Colombia and presented a key to all South American species.
APOZYGINAE
M. Sharkey
Apozyx penyai Mason
Phylogeny. In a phylogenomic analysis, Jasso-Martínez et al. (2022) recovered Apozyginae as sister to all other braconid subfamilies as suggested by Mason (1978) on morphological grounds.
Biology. The biology of A. penyai is unknown but due to its basal phylogenetic position within Braconidae and its general similarity to some members of Doryctinae it may be an idiobiont ectoparasitoid of xylophagous coleopterous larvae.
Diversity and Distribution. Apozyginae contains one rare species from Chile, Apozyx penyai Mason (Figs 3–4).
Diagnosis. (Figs 4–5). This is the only extant braconid in which forewing crossvein 2m-cu (the second recurrent vein) is present. Additionally, vein 2Cub is present in the hind wing, and the oral cavity is cyclostome.
DIRRHOPINAE
J.B. Whitfield and M. Sharkey
Dirrhope Foester
Phylogeny. Within Braconidae, Dirrhopinae is clearly a part of the microgastroid clade (Quicke and van Achterberg, 1990; Wharton et al., 1992; Whitfield and Mason, 1994). Recently Dirrhope was included in a molecular phylogenetic analysis which found it to be the sister taxon to Cheloninae, confirming the microgastroid placement (Jasso-Martínez et al. 2023).
Biology. These are koinobiont endoparasitoids of Nepticulidae (Lepidoptera) (van Achterberg 1984; Whitfield and Wagner 1991).
Diversity. Only a single genus, Dirrhope Foester, 1851 (Fig. 6) is known. There are six described species, and probably a dozen or two are undescribed. There is only one New World species described, D. americana Muesebeck, which is widespread in the USA but rare.
Distribution. Cosmopolitan, reported here from Africa and the Neotropical region for the first time (Madagascar, South Africa, and Costa Rica, all in the Hymenoptera Institute Collection).
Diagnosis. Specimens of Dirrhope resemble those of Adelius (Cheloninae), Miracinae, and small Microgastrinae in size and reduced venation. Dirrhope species differ from these taxa in their unique forewing venation (Figs 7–8).
Publications. Muesebeck (1936) described D. americana. Ranjith et al. (2021) revised and keyed the six world species.
MASONINAE
M. Sharkey, C. van Achterberg, and D.L.J. Quicke
Masona van Achterberg
Phylogeny. Quicke et al. (2019b) suggested, using multiple molecular markers, that Masoninae belongs in Ichneumonidae. Jasso-Martínez et al. (2022) recovered Masoninae as sister to Aphidiinae and we follow their placement here.
Biology. Unknown.
Diversity. Masona van Achterberg (Fig. 9) is the only extant genus, and it has seven described species. It has been collected rarely, primarily in flight-interception traps (including the wingless females) and Winkler traps.
Distribution. Two of the seven described extant species occur in southeastern USA. Two species are known from Australia. Quicke et al. (2019a) described a new species from Brazil, and Dal Pos et al. (2024) described a new species from Puerto Rico. There is also a report of an undescribed species from Tanzania (van Achterberg, 1995) and Asia (Quicke et al., 2019b). The specimen shown in Fig. 10 represents a new species from Colombia and the specimen in Fig. 9 is from California, USA.
Diagnosis. Masona is a small genus with specimens 1–2 mm long. They are sexually dimorphic. Females are wingless (Figs 10, 12) with an ant-like habitus, head elongated behind eyes, antenna distinctly shorter than body and with 14–19 segments, and the head in New World species distinctly prognathous. The winged males are mainly characterized by the short face and head and the wings are almost devoid of venation (Fig. 11). The upcurved ovipositor, the shortened foretarsus of females, and the labial palp with one segment in both sexes are diagnostic. The occipital carina is present dorsally and absent lateroventrally.
Publications. Van Achterberg (1995) erected the subfamily and genus and described four species. Two amber fossils from the New World have been described from Miocene Dominican amber, one in the extant genus Masona (van Achterberg, 2001) and one in a new, probably extinct, genus, Anoblepsis (Engel and Bennett, 2008). Dal Pos et al. (2024) revised the world species, included a key, described two new species, and transferred M. timpaynei Quicke incertae sedis to Neorhacodinae (Ichneumonidae).
METEORIDEINAE
M. Sharkey
Meteoridea Ashmead, 1900
Phylogeny. Sharanowski et al. (2011) found a sister-group relationship between Meteoridea and the Sigalphoid + Microgastroid clade. Jasso-Martínez et al. (2022) recovered Meteorideinae, represented by Meteoridea hutsoni Nixon, as sister to all other non-cyclostome braconids.
Biology. Members are known to be gregarious or solitary, larval-pupal endoparasitoids of Lepidoptera (Nixon 1941, Ghosh and Abdurahiman 1984). Eggs are laid into the host larva, and the adult parasitoid emerges from the pupal stage of the host. These generalities are based primarily on studies of Meteoridea hutsoni, a parasite of Opisina arenosella Walker, a pest of coconut. The largest collections of Meteoridea (Canadian National Collection) have been taken from light traps, suggesting that at least some species are nocturnal.
Diversity. Only Meteoridea (Fig. 13) is found in the New World. There are 16 described species worldwide, two from the Neotropics and two from the Nearctic region. There are many more undescribed species, but probably 30–70 in the New World. Yu et al. (2016) catalogued the world species of Meteoridea.
Distribution. Cosmopolitan and much more diverse in the Neotropics, as are most braconid subfamilies.
Diagnosis. Meteorideines can be recognized by the following combination of characters: hind wing with vein Cub present although often very small and difficult to see; forewing with second submarginal cell quadrate; terminal segments of metasoma laterally compressed; ovipositor very small, barely (if at all) exerted from the apex of the metasoma.
Publications. Penteado-Dias (1996) described the two Neotropical species and included a key to the four described New World species.
Acknowledgements
Thank you to José Fernandez-Triana and John Huber for their reviews, to Donna Giberson and Heather Proctor for major reformatting of the text, and to other editors of CJAI for their patience and tireless work.
References
Arias-Penna, D.C. 2007. New geographical records of the genus Urosigalphus Ashmead, 1889 (Hymenoptera: Braconidae, Helconinae) for Colombia. Boletín del Museo de Entomología de la Universidad del Valle, 8(1): 1–9.
Arias-Penna, D.C. and Whitfield, J.B.A. 2012. Taxonomic revision of the Colombian species of Urosigalphus Ashmead (Hymenoptera: Braconidae). Zootaxa, 3411: 1–54. https://doi.org/10.11646/zootaxa.3411.1
Belshaw, R. and Quicke, D.L.J. 2002. Robustness of ancestral state estimates: evolution of life history strategy in ichneumonoid parasitoids. Systematic Biology, 51: 450–477.
Dal Pos, D., Broad, G.R. and Martens, A.P. 2024. Small jewels: two new species of the rare genus Masona van Achterberg (Hymenoptera, Ichneumonoidea, Braconidae), with a catalogue of world species and comments on the peculiar morphology of the genus. European Journal of Taxonomy, 925: 135–160. https://doi.org/10.5852/ejt.2024.925.2457
Engel, M.S. and Bennett, D.J. 2008. Anoblepsis, a new, bizarre braconid wasp genus in Dominican amber (Hymenoptera: Braconidae). Journal of the Kansas Entomological Society, 81: 368–372. https://www.jstor.org/stable/25568926
Ghosh, S.M. and Abdurahiman, U.C., 1984. Bioethology of Meteoridea hutsoni (Nixon) (Hymenoptera: Braconidae), a parasite of Opisina arenosella Walker, the black headed caterpillar pest of coconut. Entomon, 9: 31–34.
Gibson, L.P. 1972a. Revision of the genus Urosigalphus of the United States and Canada (Hymenoptera: Braconidae). Miscellaneous Publications of the Entomological Society of America, 8(3):85–134. https://doi.org/10.4182/MZBP8924.8-3_4.85
Gibson, L.P. 1972b. Urosigalphus of Mexico and Central America (Hymenoptera: Braconidae). Miscellaneous Publications of the Entomological Society of America, 8(4):135–157. https://doi.org/10.4182/MZBP8924.8-3_4.135
Gibson, L.P. 1974. South American Urosigalphus (Hymenoptera: Braconidae). Miscellaneous Publications of the Entomological Society of America, 9(4):201–226.
Gibson, L.P. 1982a. New species of Urosigalphus (Hymenoptera: Braconidae) from Mexico. Proceedings of the Entomological Society of Washington, 84(1):97–101.
Gibson, L.P. 1982b. New species of Urosigalphus (Hymenoptera: Braconidae) from South America. Proceedings of the Entomological Society of Washington, 84(1):167–176.
Jasso-Martínez, J.M., Brady, S.G and Kula, R.R. 2023. Phylogenetic affinities of the non-cyclostome subfamilies Amicrocentrinae and Dirrhopinae (Hymenoptera, Braconidae) confirmed by ultraconserved element data. Journal of Hymenoptera Research, 96: 1017–1030. https://doi.org/10.3897/jhr.96.111012
Jasso-Martínez, J.M., Santos, B.F., Zaldívar-Riverón, A., Fernandez-Triana, J., Sharanowski, B.J., Richter, R., Dettman, J.R., Blaimer, B.B., Brady, S.G. and Kula, R.R. 2022. Phylogenomics of braconid wasps (Hymenoptera, Braconidae) sheds light on classification and the evolution of parasitoid life history traits. Molecular Phylogenetics and Evolution, 173: 107452. https://doi.org/10.1016/j.ympev.2022.107452
Mason, W.R.M. 1978. A new genus, species and family of Hymenoptera (Ichneumonoidea) from Chile. Proceedings of the Entomological Society of Washington, 80: 606–610.
Mason, W.R.M. 1981. The polyphyletic nature of Apanteles Foerster (Hymenoptera: Braconidae): a phylogeny and reclassification of Microgastrinae. Memoirs of the Entomological Society of Canada, 115: 1–147. https://doi.org/10.4039/entm113115fv
Mason, W.R.M. 1983. A new South African subfamily related to Cardiochilinae (Hymenoptera: Braconidae). Contributions to the American Entomological Institute, 20: 49–62.
Mason, W.R.M. 1987. Discovery of female Apozyx (Hymenoptera: Apozygidae) and comments on its taxonomic position. Proceedings of the Entomological Society of Washington, 89: 226–229.
Muesebeck, C.F.W. 1936. On two little known genera of Braconidae (Hymenoptera). Proceedings of the Entomological Society of Washington, 37: 173–177. (Note: though the journal is dated 1935, it was not published until early 1936.)
Nixon, G.E.J. 1941. New braconid parasites of Antestia lineaticollis Stal and of Sylepta derogata F. Bulletin of Entomological Research, 32:93–101.
Penteado-Dias, A.M., 1996. New Neotropical species of the genus Meteoridea Ashmead (Hymenoptera: Braconidae: Meteorideinae). Zoologische Mededelingen, 70: 197–205.
Quicke, D.L.J. and van Achterberg, C. 1990. Phylogeny of the subfamilies of the family Braconidae (Hymenoptera: Ichneumonoidea). Zoologische Verhandelingen Leiden, 258: 1–95.
Quicke, D.L.J., Austin, A.D., Fagan‐Jeffries, E.P., Hebert, P.D.N. and Butcher, B.A. 2019b. Molecular phylogeny places the enigmatic subfamily Masoninae within the Ichneumonidae, not the Braconidae. Zoological Scripta, 49: 64–71.
Quicke, D.L.J., Chaul, J.C.M. and Butcher, B.A. 2019a. First South American record of the rare ichneumonoid subfamily Masoninae van Achterberg (Hymenoptera: Ichneumonoidea: Ichneumonidae) with description of a new species from Brazil. Zootaxa, 4664: 587–593.
Ranjith, A.P., Samartsev, K.G. and Nasser, M. 2021. Discovery of the braconid subfamily Dirrhopinae van Achterberg (Hymenoptera: Ichneumonoidea) from the Indian subcontinent with the description of a new species from south India. Zootaxa, 4908(2): 251–262. https://doi.org/10.11646/zootaxa.4908.2.6
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. 1997a. Subfamily Apozyginae. In Manual of the New World genera of Braconidae (Hymenoptera). Edited by R.A. Wharton, P.M. Marsh, and M.J. Sharkey. Special Publication of the International Society of Hymenopterists. pp. 132–134
Sharkey, M.J. 1997b. Subfamily Helconinae, In Manual of the New World genera of Braconidae (Hymenoptera). Edited by R.A. Wharton, P.M. Marsh, and M.J. Sharkey. Special Publication of the International Society of Hymenopterists. pp. 260–272.
Sharkey, M.J. 1997c. Subfamily Meteorideinae, In Manual of the New World genera of Braconidae (Hymenoptera). Edited by R.A. Wharton, P.M. Marsh, and M.J. Sharkey. Special Publication of the International Society of Hymenopterists. pp. 324–325.
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
Sharkey, M.J. and Wharton, R.A. 1997. Morphology and terminology, pp.19-37. 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.
Stigenberg, J., Boring, C.A. and Ronquist, F. 2015. Phylogeny of the parasitic wasp subfamily Euphorinae (Braconidae) and evolution of its host preferences. Systematic Entomology, 40: 570–591. https://resjournals.onlinelibrary.wiley.com/doi/epdf/10.1111/syen.12122
Tobias, V.I. 1987. New taxa of Braconidae from Baltic amber (Hymenoptera). Entomologicheskoye Obozreniye, 66(4): 845–859. [Entomological Review, 67(4):18–32.]
van Achterberg, C. 1984. Essay on the phylogeny of Braconidae (Hymenoptera: Ichneumonoidea). Entomologisk Tidskrift, 105: 41–58.
van Achterberg, C. 1990. Pronkia, a new genus of the Meteorideinae (Hymenoptera: Braconidae) from New Zealand. Zoologische Mededelingen Leiden, 64: 169–175. https://repository.naturalis.nl/pub/318945/ZM1990064013.pdf
van Achterberg, C. 1995. Generic revision of the subfamily Betylobraconinae (Hymenoptera: Braconidae) and other groups with modified fore tarsus. Zoologische Verhandelingen Leiden, 298: 1–242.
van Achterberg, C. 1997. Subfamily Masoninae, In Manual of the New World Genera of Braconidae (Hymenoptera). Edited by R.A. Wharton, P.M. Marsh, and M.J. Sharkey. Special Publication of the International Society of Hymenopterists. pp. 316–318.
van Achterberg, C. 2001. The first known fossil Masoninae (Hymenoptera: Braconidae) from Miocene Dominican amber. Zoologische Mededelingen Leiden, 75: 393–396. https://repository.naturalis.nl/pub/217453/ZM75_393-396.pdf
van Achterberg, C. 2002. Revision of the genus Canalicephalus Gibson and the recognition of the Acampsohelconinae (Hymenoptera: Braconidae) as extant. Zoologische Mededelingen Leiden, 76(20):347–370. https://repository.naturalis.nl/pub/217479/
Wharton, R.A., Marsh, P.M. and Sharkey, M.J. (eds). 1997. Manual of the New World genera of Braconidae (Hymenoptera). Special Publication of the International Society of Hymenopterists, 439 pp.
Wharton, R.A., Shaw, S.R., Sharkey, M.J., Wahl, D.B., Woolley, J.B., Whitfield, J.B., Marsh, P.M. and Johnson, J.W. 1992. Phylogeny of the subfamilies of the family Braconidae (Hymenoptera: Ichneumonoidea): a reassessment. Cladistics, 8: 199–235. https://doi.org/10.1111/j.1096-0031.1992.tb00068.x
Whitfield, J.B. 1997a. Subfamily Dirrhopinae. In Manual of the New World genera of Braconidae (Hymenoptera). Edited by R.A. Wharton, P.M. Marsh, and M.J. Sharkey. Special Publication of the International Society of Hymenopterists. pp. 201–204.
Whitfield, J.B. and Mason, W.R.M. 1994. Mendesellinae, a new subfamily of braconid wasps (Hymenoptera: Braconidae) with a review of relationships within the microgastroid assemblage. Systematic Entomology, 19: 61–76. https://doi.org/10.1111/j.1365-3113.1994.tb00579.x
Whitfield, J.B. and Wagner, D.L. 1991. Annotated key to the genera of Braconidae (Hymenoptera) attacking Ieafmining Lepidoptera in the Holarctic Region. Journal of Natural History, 25: 733–754. https://doi.org/10.1080/00222939100770481
Yu, D.S.K., van Achterberg, C. and Horstmann, K. 2016. Taxapad 2016, Ichneumonoidea 2015. Database on flash-drive. www.taxapad.com, Ottawa, Ontario, Canada.
