Biodiversity Data Journal :
Taxonomic Paper
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Corresponding author: AJ Fleming (ajfleming604@gmail.com)
Academic editor: Daniel Whitmore
Received: 08 Aug 2017 | Accepted: 30 Nov 2017 | Published: 05 Dec 2017
© 2017 AJ Fleming, D. Monty Wood, M. Alex Smith, Tanya Dapkey, 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 AJ, Wood DM, Smith MA, Dapkey T, Hallwachs W, Janzen D (2017) A new species of Voria Robineau-Desvoidy (Diptera: Tachinidae) from Area de Conservación Guanacaste in northwestern Costa Rica. Biodiversity Data Journal 5: e20123. https://doi.org/10.3897/BDJ.5.e20123
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We describe a new species in the genus Voria Robineau-Desvoidy, 1830 (Diptera: Tachinidae: Voriini) from Area de Conservación Guanacaste (ACG) in northwestern Costa Rica. It was reared as part of an ongoing inventory of wild-caught caterpillars spanning a variety of moth and butterfly families (Lepidoptera). Our study provides a concise description of the new species using morphology, life history, molecular data, and photographic documentation. In addition to the new species, we provide a diagnosis of the genus as well as new data relating to host use.
The following new species of Voria is described: Voria erasmocoronadoi Fleming & Wood sp. n.
The following are proposed by Fleming & Wood as new synonyms of Voria: Xenoplagia Townsend, 1914 syn. n., Hystricovoria Townsend, 1928 syn. n., Afrovoria Curran, 1938 syn. n., and Anavoria Mesnil, 1953 syn. n., and Itavoria Townsend, 1931 syn. n.
The following new combinations are proposed as a result of the new synonymies: Voria bakeri (Townsend, 1928), comb. n. and Voria setosa (Townsend, 1914), comb. n. The authors also propose Voria pollyclari (Rocha-e-Silva, Lopes & Della Lucia, 1999), comb. n. based on the morphology of the holotype.
caterpillar, tropical, Voriini, Noctuidae, parasitoid, fly, rainforest, dry forest, cloud forest, ACG
Generally speaking, the tribe Voriini (Diptera: Tachinidae: Dexiinae) can be characterized by two main traits, namely: the obliquely angled sinusoidal hind crossvein, and the long ribbon-like phallus (
The genus Voria was originally erected for Voria latifrons Robineau-Desvoidy, 1830, which was described based on an unspecified number of specimens (males and females) collected on flowers of Heraclæum spondylium L. (Apiaceae) found growing in a prairie in Gentilly (a suburb of Paris, France).
The new species of Voria described herein is based on specimens collected during the ongoing inventory of the tri-trophic relationships between herbivores, host plants and parasitoids within the dry, rain, and cloud forests of the terrestrial portion of ACG (
This paper is part of a larger effort to describe new species reared during the ACG inventory (
All reared tachinid specimens were obtained from host caterpillars collected in ACG (
The description of the new species is complemented with a series of color photos, to illustrate some of the morphological characters described. The morphological terminology used follows
Salient features of Voria erasmocoronadoi sp. n.; a: female paratype n. DHJPAR0006953; c–d: male paratype n. DHJPAR0040781.
All caterpillars reared from the ACG inventory receive a unique voucher code in the format yy–SRNP–xxxxx. Any parasitoid emerging from a caterpillar receives the same voucher code as a record of the rearing event. If and when the parasitoid is later dealt with individually it receives a second voucher code unique to it, in the format DHJPARxxxxxxx. These voucher codes assigned to both the host and its parasitoids may be used to obtain the individual rearing record at http://janzen.bio.upenn.edu/caterpillars/database.lasso.
To date, all DHJPARxxxxxx-coded tachinids have had one leg removed for DNA barcoding at the Center for Biodiversity Genomics, University of Guelph, Ontario, 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 WH, 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 and WH 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. The date of collection is the date of eclosion of the fly, not the date of capture of the caterpillar. This is because 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, and therefore a date that is appropriate for comparison with adult-caught specimens in other museum collections. 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. Holotypes of all tachinid parasitoids collected by the ACG inventory, including that of the species newly described herein, are deposited at CNC.
CNC – Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa, Canada
ETHZ – Erdgenössische Technische Hochschule-Zentrum, Zurich, Switzerland
IZCAS – Chinese Academy of Sciences, Institute of Zoology, Beijing, China
MLPA – Museo de la Plata, Universidad Nacional de La Plata, La Plata, Argentina
MNHL – Museum d'Histoire Naturelle de la Ville de Lille, Lille, France
MRSN – Museo Regionale di Scienze Naturali di Torino [collection formerly housed at Museo di Zoologia, Istituto di Zoologia e Anatomia Comparata, Università di Torino – MZUT], Turin, Italy
MZLU – Museum of Zoology, Lund University, Lund, Sweden
NHMUK – Natural History Museum, London, United Kingdom [formerly British Museum (Natural History)]
NHRS – Naturhistoriska riksmuseet, Stockholm, Sweden
SANC – South African National Collection of Insects, Pretoria, South Africa
SDEI – Senckenberg Deustsches Entomologisches Institut, Müncheberg, Germany
SEMK – Snow Entomological Museum, University of Kansas, Lawrence, Kansas, USA
UFVB – Museu Regional de Entomologia, Universidade Federal de Viçosa, Viçosa, Minas Gerais, Brazil
USNM – National Museum of Natural History, Washington, D.C., USA (formerly the United States National Museum)
Names of undescribed host species follow a standardized, interim naming system used for taxonomic units considered as distinct species not yet formally identified or scientifically described but reliably identified by DNA barcodes. The interim names are given in the format "Eois Janzen52" or "Caviria reginaDHJ01", 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.
The standard DNA barcode region for animals (5’ cytochrome c oxidase I (CO1) gene) (
Voria Robineau-Desvoidy, 1830: 195. Type species: Voria latifrons Robineau-Desvoidy, 1830 [=Tachina ruralis Fallén, 1810], by monotypy.
Plagia Meigen, 1838: 201. Type species: Tachina verticalis Meigen, 1824 [=Tachina ruralis Fallén, 1810], by subsequent designation of
Xenoplagia Townsend, 1914: 13. Type species: Xenoplagia setosa Townsend, 1914, by original designation. Syn. n.
Hystricovoria Townsend, 1928: 395. Type species: Hystricovoria bakeri Townsend, 1928, by original designation. Syn. n.
Itavoria Townsend, 1931: 475. Type species: Itavoria aurescens Townsend, 1931, by original designation. Syn. n.
Afrovoria Curran, 1938: 5. Type species: Afrovoria munroi Curran, 1938 [=Hystricovoria bakeri Townsend, 1928], by original designation. Syn. n.
Anavoria Mesnil, 1953: 170 (as subgenus of Voria Robineau-Desvoidy, 1830). Type species: Voria (Anavoria) indica Mesnil, 1953 [=Hystricovoria bakeri Townsend, 1928], by monotypy. Syn. n.
Other species included in Voria Robineau-Desvoidy
aurifrons Townsend, 1892: 67 (Plagia). Holotype male (SEMK). Type locality: USA, Pennsylvania.
aurescens Townsend, 1931: 475 (Itavoria). Holotype male (USNM). Type locality: Brazil, São Paulo, Itaquaquecetuba. Comb. n.
bakeri Townsend, 1928: 395 (Hystricovoria). Holotype male (USNM). Type locality: Philippines, Luzon, Mt. Makiling [as “Mount Maquiling”]. Comb. n.
munroi Curran, 1938: 6 (Afrovoria). Holotype male (SANC). Type locality: South Africa, Mpumalanga, Barberton.
indica Mesnil, 1953: 170 (Voria (Anavoria)). Holotype female (NHMUK). Type locality: India, Uttarakhand, Dehra Dun.
capensis Villeneuve, 1935 (Voria): 138. Holotype male (not located, possibly lost or destroyed). Type locality: South Africa.
setosa Brauer & Bergenstamm, 1891: 409, 439 [also 1891: 105, 135] (Plagia) as “setosa Wd. litt. Cap. [Cape of Good Hope]”, nomen nudum.
micronychia Chao & Zhou, 1993: 1335 (Voria). Holotype male (IZCAS). Type locality: China, Yunnan, Zhongdian, 2400m.
operosa Robineau-Desvoidy, 1863: 827 (Voria), nomen dubium.
parva Johnson, 1919: 436 (Plagia). Syntypes, 2 females (not located, possibly lost or destroyed). Type locality: Jamaica, “Liguanea Plain, near Kingston”.
rufitorax Pazos, 1914: 1002 (Plagia), nomen nudum.
pollyclari Rocha-e-Silva, Lopes & Della Lucia, 1999: 85 (Cyrtophloeba). Holotype male (UFVB). Type locality: BRASIL, Minas Gerais, Viçosa [20°45′S e 40°51′W]. Comb. n.
ruralis Fallén, 1810: 265 (Tachina). Lectotype male (NHRS), by designation of
ambigua Fallén, 1810: 275 (Tachina). Holotype female (NHRS or MZLU). Type locality: Sweden.
verticalis Meigen, 1824: 299 (Tachina). Holotype male (MNHN). Type locality: Europe.
latifrons Robineau-Desvoidy, 1830: 196 (Voria). Syntypes, unspecified number and sex (MNHN). Type locality: France, Gentilly.
arcuata Macquart, 1834: 264 (Tachina). Holotype male (MNHL). Type locality: France, Lille.
interrupta Zetterstedt, 1844: 1020 (Tachina). Holotype female (NHRS). Type locality: Sweden, Scania.
transversa Macquart, 1848: 96 (Plagia). Holotype female (ETHZ). Type locality: Switzerland, Zurich.
spinicosta Palm, 1876: 419 (Tachina). Holotype female (not located, possibly lost or destroyed). Type locality: Austria, Innsbruck.
americana van der Wulp, 1890: 102 (Plagia). Syntypes, males and females (NHMUK). Type localities: Mexico: Veracruz (Orizaba); Guerrero (Venta del Zopilote); Xucumanatlan; Omilteme and Tabasco (Teapa).
mexicana Giglio-Tos, 1893: 5 (Plagia). Holotype female (MRSN). Type locality: Mexico.
brasiliana Townsend, 1929: 380 (Voria). Syntypes, males and females (USNM). Type locality: Brazil, São Paulo, Itaquaquecetuba.
edentata Baranov, 1932: 83 (Voria). Holotype male (SDEI) (see
ayerzai Blanchard, 1943: 157 (Plagia). Syntypes, unspecified number and sex (MLPA). Type locality: not given [listed as Argentina according to
ciliata d’Aguilar, 1957: 261 (as ssp. of Voria ruralis ). Holotype male (USNM). Type locality: China, Sichuan, Suifu.
saginata Walker, 1861: 298 (Eurigaster). Holotype female (NHMUK). Type locality: Mexico.
signata. Incorrect subsequent spelling of saginata Walker, 1861 (
setosa Townsend, 1914: 14 (Xenoplagia). Holotype female (USNM). Type locality: Peru, Cañada de Saman, Rio Chira, Peru. Comb. n.
Voria can be distinguished by the following combination of traits: compound eye bare; vertex, at its narrowest point, approximately 0.6X eye width in dorsal view; frontal vitta widened posteriorly, ranging in color from dark gray to gold; both sexes with well-developed lateral vertical setae; fronto-orbital plate with 2–4 proclinate orbital setae; fronto-orbital plate ranging from bare to haired; frontal setae descending below base of pedicel; parafacial with some fine hairs on upper 1/3, sometimes with a small tuft of hairs at level of vibrissa; parafacial with 1–3 strong proclinate setae below lowest frontal seta; occiput without black setae posterior to postocular row; genal dilation very slightly developed; 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; anepimeral setae not or weakly differentiated from other hairs on anepimeron; wing cell r4+5 open at wing margin; bend of vein M with a long stub; wing vein R1 setulose along its entire length; R4+5 with setulae extending from the fork to just beyond crossvein R-M; abdomen narrow, oval, black in ground color; tergites 3–5 without median discal setae.
Voria is a widespread cosmopolitan genus originally described from Sweden. The genus ranges across the Holarctic, Afrotropical, Australasian, Oriental and Neotropical regions.
Voria has been known to parasitize lepidopteran larvae primarily in the family Noctuidae (and possibly also Pyralidae) (
Male (Fig.
Female (Fig.
Voria erasmocoronadoi sp. n. can be differentiated from its congeners by the following combination of the traits: vertex, at its narrowest point, approximately 0.67X eye width in dorsal view; frontal vitta widened posteriorly, with strong gold pollinosity; fronto-orbital plate gold pollinose, with three proclinate orbital setae; frontal setae descending below base of pedicel, with sparse black hairs intermingled among setae; fronto-orbital hairs extending into upper third of parafacial; and parafacial with only one proclinate seta, below lowest frontal seta. Voria ruralis can be differentiated from V. erasmocoronadoi by the lack of gold pollinosity on the fronto-orbital plate, and the regular length and spacing of the setae on the anterodorsal surface of the fore tibia. In addition to the barcode and the external morphological character differences, our results also suggest that there are strong differences in the shapes of the postgonite and surstylus between the various populations of V. ruralis previously described and our new species. We consider a detailed comparison to exceed the scope of the present paper. Voria aurifrons can be distinguished by the presence of a slight infuscation along the costal margin of the wing, and by having legs of a reddish ground color, two character states lacking in V. erasmocoronadoi.
Voria erasmocoronadoi sp. n. is named in honor of Mr. Erasmo Coronado Caballo of Liberia, Costa Rica, in recognition of his years of dedicated logistic support to the Guanacaste Dry Forest Conservation Fund and to the ACG parataxonomist program, participants of which found and reared the caterpillar hosts of this fly.
Costa Rica, ACG (Guanacaste and Alajuela provinces), 121–1150m.
Voria erasmocoronadoi sp. n. has been reared seven times at ACG: six times from caterpillars of three species of Noctuidae, Ctenoplusia oxygramma (Geyer, 1832), Chrysodeixis includens (Walker, 1858) and Diastema morata Schaus, 1894, and once from a caterpillar of Erosia biolepDHJ03 in the family Uraniidae. Sites of collection include cloud forest, rainforest, and dry-rain intergrade forest.
The DNA barcode region sequences for Voria are AT-biased (70%), as expected for insect DNA. The sequences displayed no heteroplasmy or stop codons that would suggest the amplification of a pseudogene and we conclude that these barcodes are mtDNA for Voria. There are currently five other species in BOLD provisionally identified as Voria ruralis from localities other than ACG in BOLD, each with their own unique Barcode Index Numbers (BINs); all are likely to be members of the Voria ruralis species complex. The sequence variation between V. erasmocoronadoi sp. n. (BOLD:AAG9377) and the other Voria BINS on BOLD ranges from 3 to 11%. Based on our experience with other species complexes of morphologically indistinct insect species groups in ACG, this distance is highly supportive of the status of V. erasmocoronadoi sp. n. as a separate species.
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). This study has been supported by the Government of Canada through its ongoing support of the Canadian National Collection, Genome Canada, the Center for Biodiversity Genomics, and the Ontario Genomics Institute (2008–0GI–ICI–03) (MAS), and by a Discovery Grant from the Natural Sciences and Engineering Research Council of Canada (MAS).