Biodiversity Data Journal :
Taxonomic Paper
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Corresponding author: AJ Fleming (ajfleming604@gmail.com)
Academic editor: Torsten Dikow
Received: 12 May 2021 | Accepted: 16 Jul 2021 | Published: 29 Jul 2021
© 2021 AJ Fleming, D. Monty Wood, M. Alex Smith, Winnie Hallwachs, Daniel Janzen
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Fleming A, Wood DM, Smith MA, Hallwachs W, Janzen D (2021) Revison of Metaplagia Coquillett (Diptera: Tachinidae) with description of five new species from Area de Conservación Guanacaste in northwestern Costa Rica. Biodiversity Data Journal 9: e68598. https://doi.org/10.3897/BDJ.9.e68598
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We revise the genus Metaplagia Coquillett, 1895 and describe five new species from Area de Conservación Guanacaste (ACG) in northwestern Costa Rica. All new species were reared from an ongoing inventory of wild-caught caterpillars spanning a variety of species within the family Sphingidae (Lepidoptera: Sphingidae). Our study provides a concise description of each new species using morphology, life history, molecular data and photographic documentation. In addition to the new species, the authors provide a re-description of the genus and a revised key to the species of Metaplagia.
The following five new species of Metaplagia are described: Metaplagia leahdennisae Fleming & Wood sp. n., Metaplagia lindarobinsonae Fleming & Wood sp. n., Metaplagia paulinesaribasae Fleming & Wood sp. n., Metaplagia robinsherwoodae Fleming & Wood sp. n. and Metaplagia svetlanakozikae Fleming & Wood sp. n.
The following is proposed by Fleming & Wood as new combination of Plagiomima Brauer & Bergenstamm, 1891: Plagiomima latifrons (Reinhard, 1956) comb. n.
tropical rain forest, tropical dry forest, cloud forest, parasitoid flies, host-specificity, caterpillars, ACG, Dexiinae, Voriini
The tribe Voriini is distributed globally; the vast tachinid fauna of the Neotropical Region and the huge number of genera have proven to be one of the most significant hurdles to understanding the tribal boundaries surrounding the Voriini. Most attempts to understand the synapomorphies or of the tribe have, rather than answered questions, led researchers to even more questions as to what the voriines truly are. Presently, the tribe possesses several diagnostic features which the authors consider as typical to the tribe: conical head profile (longer at level of pedicel than at vibrissa); proclinate, divergent and well-developed ocellar setae; frons wide; proclinate and reclinate orbital setae present in both sexes; facial ridge bare; prosternum bare; anepimeral seta absent or poorly developed appearing hair-like; infrasquamal setae present; apical scutellar setae strong and decussate; dm-m crossvein oblique, making posterior section of M4 subequal to anterior section; R4+5 setulose at least to crossvein r-m and sometimes beyond; middorsal depression of ST1+2 reaching posterior margin; and aedeagus elongate and frequently ribbon-like (
The monotypic genus Metaplagia Coquillett, 1895 (Dexiinae: Voriini) was initially erected for the type species Metaplagia occidentalis Coquillett, 1895 (
Our goal is to systematically revise and analyze the known members of the New World tachinid genus Metaplagia Coquillett and add to the existing taxa five new species from Area de Conservación Guanacaste (ACG). Our species concepts are based on differences in morphology, life history and by comparison of COI (coxI or cytochrome c oxidase I) gene sequences. We also provide a key to the species of Metaplagia inclusive of the Nearctic and Mesoamerican Regions. This paper is part of a broader effort to name and catalog all of the tachinid species collected from the ACG inventory, an effort which has already presented new information within the tribe Voriini (
All reared specimens were obtained from host caterpillars collected in ACG (
The management of voucher specimens has been detailed in previous papers in this series (
To date, all DHJPARxxxxxx-coded tachinids have had one leg removed for DNA barcoding at the Biodiversity Institute of Ontario (BIO) in Guelph, ON, Canada. All successful barcodes and collateral data are first deposited in the Barcode of Life Data System (BOLD, www.boldsystems.org) (
Inventoried Tachinidae were collected under Costa Rican government research permits issued to DHJ and exported from Costa Rica to Philadelphia, en route to their final depository in the Canadian National Insect collection in Ottawa, Canada (CNC). Tachinid identifications for the inventory were done by DHJ in coordination with: a) visual inspection by AJF and DMW, b) DNA barcode sequence examination by MAS and DHJ and c) correlation with host caterpillar identifications by DHJ and WH through the inventory itself. Dates of collection, cited for each ACG specimen, are the dates of eclosion of the fly, not the date of capture of the caterpillar since the fly eclosion date is much more representative of the time when that fly species is on the wing than is the time of capture of the host caterpillar. The collector listed on the label is the parataxonomist who found the caterpillar, rather than the person who retrieved the newly-eclosed fly from its rearing container. The holotypes and paratypes of the species newly-described herein are all deposited at CNC.
Species accounts and descriptions are deliberately brief and concise, complemented by a series of color photos of every species, used to illustrate the morphological differences amongst them. The morphological terminology used follows
Names of undescribed host species follow a standardized, interim naming system used for taxonomic units considered as distinct species and identified by DNA barcodes. The interim names are given in the format "Manduca sextaDHJ02" or "Manduca sextaDHJ03", where the "species epithet" is either composed of the name of the taxonomist who identified the species and a number or the name of a species-group, followed by a code. This prevents confusion with already-described species, while maintaining traceability of each undescribed species within the ACG project.
We generated DNA extracts from single legs using a glass fibre protocol (
The phylogeny (Fig.
CNC, Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa, Canada
UCDC, University of California, Davis, Bohart Museum of Entomology, Davis, California, USA
USNM, United States National Museum of Natural History, Washington, D.C., USA
Metaplagia Coquillett, 1895: 102 [also 1895: 97]. Type species: Metaplagia occidentalis Coquillett, 1895, by original designation.
Metavoria Townsend, 1915: 101. Type species: Metavoria orientalis Townsend, 1915, by original designation. Synonymy proposed by
Agathomyia Reinhard, 1959: 228 (preocc. by Verrall, 1901). Type species: Agathomyia cordata Reinhard, 1959, by original designation. Synonymy proposed by
Anzamyia Reinhard, 1960: 103. (nomen novum for Agathomyia, 1959 Reinhard).
Other species included in Metaplagia Coquillett
Metaplagia brevicornis Brooks, 1945: 81. Holotype male (CNC), by original designation. Type locality: Canada, Manitoba, Teulon.
Agathomyia cordata Reinhard, 1959: 229. Holotype female (UCDC) (1 female paratype in CNC), by original designation. Type locality: USA, California, Riverside County, Anza.
Metavoria facialis Reinhard, 1956: 123. Holotype female (CNC). Type locality: USA, Utah, Fruitland.
Metaplagia occidentalis Coquillett, 1895: 103. Holotype male (USNM, Type No. USNMENT01519725), by original designation. Type locality: USA, California, San Diego County.
Metavoria orientalis Townsend, 1915: 101. Holotype female (USNM, Type No. USNMENT01519726), by original designation. Type locality: USA, Virginia, Arlington.
Male, head: frontal vitta wide, 1/3-1/6th width of front-orbital plate; with 2–3 proclinate orbital setae and 1–2 reclinate orbital setae; ocellar setae proclinate slightly divergent; eye bare, not descending beyond the level of the vibrissa; fronto-orbital plate coloration ranging from shining silver to gold; fronto-orbital plate with short black setulae interspersed amongst frontal setae; fronto-orbital setae not extending below lower margin of pedicel, with fronto-orbital setulae sometimes extending below lower margin of pedicel; pedicel orange, with a black-dark brown post pedicel; arista bare and subequal to slightly shorter than postpedicel, distinctly-thickened on basal 1/2-2/3, ranging from dark orange to dark brown-black; lower margin of face almost level with vibrissa, not visible in profile; facial ridge bare, but with setulae along parafacial sometimes so close to facial ridge as to be confused with facial ridge setulae; palps either straight or with a slight club at apex, sparsely setulose. Thorax: tomentosity ranging from pale beige-gray to dark grey or silver over a black ground color; thorax black ground color tomentum of thorax ranging from pale brassy to silver grey; prosternum bare; 3–4 postpronotal setae arranged in a straight line; supra-alar setae 1–2:3; intra-alar setae 2–3:3; dorsocentral setae 3:3–4; acrostichal setae 3:3; katepisternum with 3 setae. Scutellum black ground color, with tomentum ranging from gray to pale brassy; with five pairs marginal setae; apical scutellar setae 1/2–1/3 as long as subapical scutellars, sub-erect, slightly above marginal plane; basal scutellar subequal in length to subapical setae; subapical setae straight, ranging from divergent to convergent. Legs: dark reddish-brown to black ground color; tarsal claws and pulvilli ranging from shorter than to longer than last tarsomere. Wings: slightly longer than abdomen; translucent slightly hyaline; R1 and R4+5 can be setulose, setulae of R4+5 ranging from node to crossvein r-m or beyond. Abdomen: ground color black, pale silver tomentum in varying degrees on T3–T5, ST1+2 typically glabrous; middorsal depression on ST1+2 reaching to hind margin of tergite; median marginal setae present on T3 and complete rows on T4 and T5; median discal setae absent on all tergites; sex patch absent.
Terminalia: posterior margin of sternite 5 with a deeply excavated and wide U-shaped (sometimes sculptured) median cleft; lateral lobes of sternite apically rounded, often with setae along caudal margin; basal section 1/5 the length of apical lobes. Epandrium often with 3–4 long, strong setae along anal edge. Cerci, in posterior view, medially separated, but parallel and often touching so as to appear fused, with a few short setae on basal half. In lateral view, bowed and sharply tapered apically. Surstylus well-developed, stout basally in lateral view, like a stout broadly rounded triangle terminating in a small knob, appearing hooked or slightly beaked apically; in posterior view, basally enlarged and apically straight.
Female as in male, except in the following aspects: head: tomentum of fronto-orbital plate and parafacial can sometimes differ from that of the conspecific male; fronto-orbital plate and parafacial up to 0.5x wider than in males.
Metaplagia can be distinguished by the following combination of traits: head distinctly conical; males with upper frontal setae reclinate; proclinate orbital setae in both sexes; frontal setae descending below level of pedicel; both sexes with well-developed lateral vertical setae; eye bare; parafacial setulose, but not with strong stout setae, only hair-like setulae; genal dilation very slightly developed; prementum shorter than height of head with an enlarged labellum; prosternum bare; three postsutural supra-alar setae, the anteriormost reduced and much weaker than first postsutural dorsocentral seta; scutellum with four pairs of marginal setae and one pair of erect to semi-erect apical setae; one or two pairs of sub-erect discal setae on scutellum, in line with subapical setae; vein M1 ending separately in wing margin; anepimeral seta short, not extending beyond edge of lower calypter; wing vein R4+5 setulose.
From Manitoba, Canada east to Newfoundland and south to Costa Rica.
Within the ACG inventory, Metaplagia has only been reared from the Lepidoptera family Sphingidae: Lepidoptera throughout the diverse ecosystems of the research area.
Our present revision of Metaplagia increases the range of the genus, taking it from being a Nearctic endemic genus to a complete New World distribution, inclusive of both the Nearctic and Neotropical Regions.
Both Plagiomima Brauer & Bergenstamm, 1891 and Metaplagia are extremely similar genera; however, these can be very easily distinguished by the conformation of their mouthparts (
Male (Fig.
Terminalia (Fig.
Female (Fig.
Metaplagia leahdennisae sp. n. can be distinguished from all other Metaplagia by the following combination of traits: fronto-orbital plate at least 50% gold, with a silver parafacial and setulae on vein R4+5 not extending beyond crossvein r-m. Metaplagia leahdennisae sp. n. can be separated from Metaplagia occidentalis by the presence of gold on the fronto-orbital plate. Metaplagia leahdennisae sp. n. is clearly distinguished by its COI sequence clustered within the Barcode Identification Number (BIN) BOLD:AAE2876.
Metaplagia leahdennisae sp. n. is named in honor of Leah Dennis for her many years of coordinating and administrating a plethora of problems and events in the Academic Office of the Department of Biology, University of Pennsylvania, Philadelphia, Pennsylvania.
Costa Rica, ACG, Guanacaste Province, 5–275 m elevation.
Metaplagia leahdennisae sp. n. has been reared eight times from one species of Lepidoptera in the family Sphingidae: Agrius cingulata (Fabricius, 1775) in dry forest.
Male (Fig.
Terminalia (Fig.
Female (Fig.
Metaplagia lindarobinsonae sp. n. can be distinguished from all other Metaplagia by the following combination of traits: frontal vitta indistinct around ocellar triangle, infrasquamal setae present, fronto-orbital plate at pale gray or silver, with a silver parafacial and setulae on vein R4+5 not extending beyond crossvein r-m, surstylus sharp at tip. Metaplagia lindarobinsonae sp. n. can be separated from Metaplagia occidentalis by the presence of infrasquamal setae and from Metaplagia svetlanakozikae sp. n. by the presence of a distinct row of setulae along the facial ridge. Metaplagia lindarobinsonae sp. n. is clearly distinguished by its COI sequence clustered within the Barcode Identification Number (BIN) BOLD:AAD5563.
Metaplagia lindarobinsonae sp. n. is named in honor of Linda Robinson for her many years of coordinating and administrating a plethora of problems and events for the undergraduate biology teaching laboratories of the Department of Biology, University of Pennsylvania, Philadelphia, Pennsylvania.
Costa Rica, ACG, Guanacaste Province, 85–300 m elevation.
Metaplagia lindarobinsonae sp. n. has been reared 17 times from two species of Lepidoptera in the family Sphingidae: Manduca sexta (Linnaeus, 1763), Manduca sextaDHJ02 and Manduca sextaDHJ03, in dry forest.
Male (Fig.
Terminalia: holotype male not dissected.
Female: unknown at this time.
Metaplagia paulinesaribasae sp. n. can be easily distinguished from all other Metaplagia by the following combination of traits: both fronto-orbital and parafacial gold and setulae on vein R4+5 not extending beyond crossvein r-m. It is distinguishable from all other congeners by the presence of an entirely gold fronto-orbital plate and parafacial. Metaplagia paulinesaribasae sp. n. is clearly distinguished by its COI sequence clustered within the Barcode Identification Number (BIN) BOLD:AAX4230.
Metaplagia paulinesaribasae sp. n. is named in honor of Pauline Saribas for her many years of coordinating and administrating a plethora of problems and events in the Academic Office of the Department of Biology, University of Pennsylvania, Philadelphia, Pennsylvania.
Costa Rica, ACG, Guanacaste Province, 280 m elevation.
Metaplagia paulinesaribasae sp. n. has been reared once from one species of Lepidoptera in the family Sphingidae: Manduca lefeburii (Guérin-Méneville, 1844), in dry forest.
Male (Fig.
Terminalia (Fig.
Female (Fig.
Metaplagia robinsherwoodae sp. n. can be distinguished from all other Metaplagia by the following combination of traits: fronto-orbital plate at least 75% gold, with a silver parafacial and setulae on vein R4+5 extending well beyond crossvein r-m. Metaplagia robinsherwoodae sp. n. can be separated from M. leahdennisae and M. paulinesaribasae by the presence of setulae on R4+5 extending well beyond crossvein r-m and it is separated from M. facialis and M. cordata by the presence of gold on its fronto-orbital plate. Metaplagia robinsherwoodae sp. n. is clearly distinguished by its COI sequence clustered within the Barcode Identification Number (BIN) BOLD:AAB3286.
Metaplagia robinsherwoodae sp. n. is named in honor of Robin Sherwood for her many years of coordinating and administrating a plethora of problems and events in the Academic Office of the Department of Biology, University of Pennsylvania, Philadelphia, Pennsylvania.
Costa Rica, ACG, Guanacaste Province, 250–300 m elevation.
Metaplagia robinsherwoodae sp. n. has been reared 28 times from one species of Lepidoptera in the family Sphingidae: Manduca rustica (Fabricius, 1775), in dry forest.
Male (Fig.
Terminalia (Fig.
Female (Fig.
Metaplagia svetlanakozikae sp. n. can be distinguished from all other Metaplagia by the following combination of traits: frontal vitta indistinct around ocellar triangle, infrasquamal setae present, fronto-orbital plate at pale gray or silver, with a silver parafacial and setulae on vein R4+5 not extending beyond crossvein r-m, surstylus rounded at tip. Metaplagia svetlanakozikae sp. n. can be separated from Metaplagia occidentalis by the presence of infrasquamal setae and from Metaplagia lindarobinsonae sp. n. by the presence of a small circular formation of setulae on the parafacial and not distinct row of setulae along the facial ridge. Metaplagia svetlanakozikae sp. n. is clearly distinguished by its COI sequence clustered within the Barcode Identification Number (BIN) BOLD:AAD5456.
Metaplagia svetlanakozikae sp. n. is named in honor of Svetlana Kozik for her years of coordinating and administrating a plethora of problems and events for the undergraduate biology teaching laboratories of the Department of Biology, University of Pennsylvania, Philadelphia, Pennsylvania.
Costa Rica, ACG, Guanacaste Province, 10–280 m elevation.
Metaplagia svetlanakozikae sp. n. has been reared seven times from one species of Lepidoptera in the family Sphingidae: Agrius cingulata, in dry forest.
Key to the Metaplagia of North and Mesoamerica |
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1 | Setulae present on wing vein R4+5 not extending beyond crossvein r-m | 2 |
– | Setulae present on wing vein R4+5 extending beyond crossvein r-m | 8 |
2 | Fronto-orbital plate and parafacial entirely and uniformly gold | paulinesaribasae sp. n. |
– | Fronto-orbital plate at most with some gold present and parafacial not gold | 3 |
3 | Fronto-orbital plate bearing some gold | leahdennisae sp. n. |
– | Fronto-orbital plate pale grey or silver (can include pale brassy) distinctly not gold | 4 |
4 | Infrasquamal setae absent | occidentalis Coquillett, 1895 |
– | Infrasquamal setae present | 5 |
5 | Frontal vitta indistinct around ocellar triangle, not reaching occiput | 6 |
– | Frontal vitta wide and prominent extending around ocellar triangle to occiput | 7 |
6 | Parafacial setulae descending along parafacial in a row near facial ridge; surstylus sharp at tip | lindarobinsonae sp. n. |
– | Parafacial setulae present as a small circular grouping along upper 1/3 of parafacial, not in a row near facial ridge; surstylus rounded at tip | svetlanakozikae sp. n. |
7 | Postpedicel just over 3 times as long as pedicel; claws and pulvilli shorter than last tarsomere | brevicornis Brooks, 1945 |
– | Postpedicel 2 times as long as pedicel; claws and pulvilli longer than last tarsomere | orientalis Townsend, 1915 |
8 | Fronto-orbital plate gold on at least upper two thirds | robinsherwoodae sp. n. |
– | Fronto-orbital plate entirely silver | 9 |
9 | Wing vein R1 bare throughout | facialis Reinhard, 1956 |
– | Wing vein R1 setulose throughout | cordata Reinhard, 1960 |
We gratefully acknowledge the unflagging support of the team of ACG parataxonomists (Janzen et al. 2009, Janzen & Hallwachs 2011) who found and reared the specimens used in this study and the team of biodiversity managers who protect and manage the ACG forests that host these tachinids and their caterpillar hosts. The study has been supported by U.S. National Science Foundation grants BSR 9024770 and DEB 9306296, 9400829, 9705072, 0072730, 0515699 and grants from the Wege Foundation, International Conservation Fund of Canada, Jessie B. Cox Charitable Trust, Blue Moon Fund, Guanacaste Dry Forest Conservation Fund, Area de Conservación Guanacaste, Permian Global and University of Pennsylvania (DHJ & WH) and in-kind support from INBio and the Museo Nacional de Costa Rica. This study has also been supported by the Government of Canada through its ongoing support of the Canadian National Collection, Genome Canada, the Biodiversity Institute of Ontario and the Ontario Genomics Institute (2008–0GI–ICI–03) (MAS) and by a Discovery Grant from Natural Sciences and Engineering Research Council of Canada (MAS). AJF and DMW would also like to extend a very special thank you Dr. Torsten Dikow (USNM), Dr. Nigel Wyatt (BMNH), Dr. David Grimaldi (AMNH) and his technician Courtney Richenbacher who provided invaluable information relative to types in their museums. Acknowledgements to Dr. James E. O'Hara and Shannon Mahony Henderson, for allowing us to make use of their indispensable literature library and World Tachinidae Database (MW & AJF).