Biodiversity Data Journal :
Taxonomic Paper
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Corresponding author:
Academic editor: Daniel Whitmore
Received: 02 Dec 2015 | Accepted: 15 Apr 2016 | Published: 19 Apr 2016
© 2016 AJ Fleming, D. Monty Wood, M. Alex Smith, Winnie Hallwachs, Daniel Janzen, Tanya Dapkey
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 D, Smith M, Hallwachs W, Janzen D, Dapkey T (2016) Two new species of Erythromelana Townsend, 1919 (Diptera: Tachinidae) from Area de Conservación Guanacaste in northwestern Costa Rica. Biodiversity Data Journal 4: e7386. https://doi.org/10.3897/BDJ.4.e7386
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We describe two new species in the genus Erythromelana Townsend, 1919 from Area de Conservación Guanacaste (ACG) in northwestern Costa Rica. Both species were reared from wild-caughtcaterpillars of Eois spp. (Lepidoptera: Geometridae). We provide a concise description of each species using morphology, life history, molecular data, and photographic documentation.
Erythromelana jimmychevezi Fleming & Wood sp. nov.
Erythromelana glenriverai Fleming & Wood sp. nov.
tropical rain forest, tropical dry forest, cloud forest, parasitoid flies, host-specificity, caterpillars, ACG, Exoristinae, Blondeliini
The tachinid genus Erythromelana Townsend, 1919 (Exoristinae: Blondeliini) is a small Neotropical genus in the tribe Blondeliini, occurring from southern Mexico to Bolivia and Brazil. Townsend originally described the genus based on one male and one female collected in Jaen province (Peru) and described as E. jaena Townsend. The genus remained untouched until
In 2013, an in-depth analysis and phylogeny of the genus Erythromelana was provided by
CNC - Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa, Canada
All flies and rearing information described here were obtained from the 35+ year–old ongoing inventory of the caterpillars, their food plants and their parasitoids of the dry forest, rain forest, cloud forest and intergrades of the 125,000+ ha terrestrial portion of Area de Conservación Guanacaste (ACG) in northwestern Costa Rica (
Descriptions of new species discussed in this paper are deliberately brief, and only include characters commonly used in tachinid fly identification. The descriptions are complemented with color photos, in order to illustrate the readily observed inter-specific differences.
Photographs of the habitus and terminalia were taken using the methods outlined in Fleming et al. (2014a). In brief, raw image files were first processed with Adobe Photoshop CS6, and then digitally stacked to produce a final composite image using the Zerene Stacker software v1.04. Imaging of the male terminalia followed methods described in Fleming et al. 2014a. Dissections of the terminalia followed the methods described in
The morphological terminology and measurements of body parts follow Inclan and Stireman (2013).
Wherever a specimen label has been examined, the information is presented using "/" to indicate the end of a label and the beginning of the next. Labels are presented from the top-most (closest to the specimen) to the bottom-most, with any additional comments given in square brackets" []".
DNA barcodes (the standard 5’ region of the mitochondrial cytochrome oxidase 1 (CO1) gene) for all ACG inventory specimens were obtained using DNA extractions made from single legs, using a glass fiber protocol (
All caterpillars collected from the ACG efforts receive a unique voucher code in the format yy–SRNP–xxxxx. Any parasitoid emerging from a caterpillar receives the same voucher code, if/and when it is individually processed for DNA barcoding, it then receives a second voucher code unique to it, in the format DHJPARxxxxxxx. These voucher codes, assigned to the host and to any emerging parasitoids, can be looked up at http://janzen.bio.upenn.edu/caterpillars/database.lasso. To date, all voucher-coded tachinids have had one leg removed for attempted DNA barcoding at the Biodiversity Institute of Ontario (BIO) in Guelph, with all collateral data and all successful barcodes permanently and publicly deposited in the Barcode of Life Data System (BOLD, www.boldsystems.org) (Ratnasingham and Hebert 2007), and later migrated to GenBank as well. The inventory is dynamic and continually growing, with regular additions of new specimens. Erythromelana sequences can be found by searching for "Erythromelana" in BOLD. Each barcoded specimen has been assigned accession codes from the Barcode of Life Data System (BOLD) and GenBank.
The inventoried Tachinidae were collected under Costa Rican government research permits issued to DHJ and likewise exported under permit by DHJ from Costa Rica to Philadelphia, and then to their final depository in the CNC. Tachinid identifications for the inventory were done by DHJ in coordination with a) visual inspection by AJF and DMW, b) DNA barcoding by MAS, and c) correlation with host caterpillar identifications by DHJ and WH through the ACG inventory itself. Dates of capture of each specimen are the dates of eclosion of the flies, and not the dates of capture of the caterpillars since the 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 caterpillar. The collector listed is the parataxonomist who collected the caterpillar, rather than the person who retrieved the newly eclosed fly and processed it by freezing, pinning, labeling and oven–drying. The biologies and parasitization rates of the flies will be the subject of later papers.
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 "Eois Janzen52", where the species epithet is composed of the name of the taxonomist who identified the species and a number. This prevents confusion with already described species while maintaining traceability of each undescribed species within the ACG project.
Erythromelana Townsend, 1919
Erythromelana Townsend, 1919a: 174. Type species: Erythromelana jaena Townsend, 1919a, by original designation. Other references: Guimarães (1971: 106); Wood (1985: 39–40); Wood & Zumbado (2011: 1403).
Minthomyia Townsend, 1919b: 564. Type species: Minthomyia abdominalis Townsend, 1919b, by original designation. Other references: Guimarães (1971: 41); Wood (1985: 39–40) (as synonym of Erythromelana).
Erythromelana can be distinguished from other Blondeliiniby the following combination of characters: Head: proclinate orbital setae absent in male, female with 2 setae; 1–2 pairs of reclinate outer orbital setae, lowermost outer orbital seta distinctly longer than uppermost frontal seta; ocellar setae ranging from absent to well developed; eyes haired, of variable density in different species; parafacial bare and extremely narrow, at narrowest point equal to or narrower than the basal width of the palpus; parafacial light gray in ground color, covered with silvery–gold pollinosity, and bare; fronto-orbital plate and vertex black in ground color, covered with a dull silver-gold pollinosity (appearing mostly black (Fig. 2b)), and with faint golden reflections visible only in lateral view (mostly on vertex); lower margin of face level with vibrissa; vibrissa positioned at extreme anteroventral corner of face; facial ridge with a few short hairs on lower third or less; arista black on basal 1/3–1/4, becoming light brownish to orange distally, elongate, and minutely pubescent, thickened only on basal 1/4; palpus light orange to yellow, sometimes darkened basally. Thorax: ground color shiny black; scutum silver pollinose presuturally, postsuturally a polished black in ground color; prosternum setose; postpronotum bearing 2 or, rarely, 3 setae; katepisternum with 2–3 setae; first postsutural supra-alar seta small or sometimes absent; apical scutellar setae lacking; wingvein R4+5 setulose at base and vein R1 setulose or bare; vein M smoothly curved at bend and ending at wing margin anterior to wing tip, separately from vein R4+5; legs ranging from entirely yellow to entirely black. Abdomen: ground color ranging from yellow-orange to black, without strong banding; mid-dorsal depression not extending to hind margin of T1+2; discal setae only present on T5. Male terminalia: sternite 5 with median cleft either U- or V-shaped; inner margin bearing minute setae; apical lobe rounded or pointed apically with either a single, long, well-developed seta, multiple well developed setae, or bare; pregonite curved anteriorly, with strong setae along posterior margin; postgonite short with curved apex; epiphallus small, usually difficult to see between the pregonites; surstylus with setulae on inner and outer surfaces, or completely bare; surstylus, in lateral view, varied in shape from almost straight to slightly concave on anterior or posterior margins, usually with a broad, rounded apex, occasionally truncate; surstylus and cercus usually subequal in length, sometimes cercus shorter than surstylus; in posterior view, cerci narrowed on apical 1/3.
Based on synapomorphieswithin the male terminalia of Erythromelana described by
arciforceps Inclan, 2013: 35. Holotype male (CNC). Type locality: Brazil, S.C., Nova Teutonia. Type label: Nova Teutonia S. C.-Brazil Nov. 1970 F. Plaumann/ HOLOTYPE Erythromelana arciforceps Inclan D.J/ DI412CA. Additional specimens examined: Brazil. [Examined by AJF.]
catarina Inclan, 2013: 32. Holotype male (CNC). Type locality: Brazil, S.C., Nova Teutonia. Type label: Nova Teutonia S. C.-Brazil Jun. 1970 F. Plaumann/ HOLOTYPE Erythromelana catarina Inclan D.J/ DI392CA. Additional specimens examined: Brazil. [Examined by AJF.]
convexiforceps Inclan, 2013: 33. Holotype male (CNC). Type locality: Mexico, Oaxaca, Suchistepec. Type label: Mexico, Oax 4.6 km Suchistepec 23.VII.1992 D.M. Wood 2150m/ HOLOTYPE Erythromelana convexiforceps Inclan D.J./ DI54CA. [Examined by AJF.]
cryptica Inclan, 2013: 29. Holotype male (CNC). Type locality: Venezuela, Aragua, Rancho Grande. Type label: VENEZUELA Aragua Rancho Grande 18-27.II.1971 G.&M. Wood 1100m/ HOLOTYPE Erythromelana cryptica Inclan D.J./ DI477CA. Additional specimens examined: Venezuela, Bolivia, Mexico and Ecuador. [Examined by AJF.]
distincta Inclan, 2013: 39. Holotype male (CNC). Type locality: Venezuela, Aragua, Rancho Grande. Type label: VENEZUELA Aragua 11 km Rancho Grande 25.II.1971 G.&M. Wood/ HOLOTYPE Erythromelana distincta Inclan D.J./ DI280CA. Additional specimens examined: Brazil and Venezuela. [Examined by AJF.]
napensis Inclan, 2013: 37. Holotype male (CNC). Type locality: Ecuador, Napo prov., Yanayacu Biological Station. Type label: ECUADOR: Napo Prov. Yanayacu Biological Station S00°35.9', W77°53.4', 2163 m REARED October 2005 8135/ HOLOTYPE Erythromelana napensis Inclan D.J. [Examined by AJF.]
woodi Inclan, 2013: 42. Holotype male (CNC). Type locality: Costa Rica, Puntarenas, Monteverde. Type label: COSTA RICA Pnts Monteverde 28.VIII.1993 D.M. Wood 1842m/ HOLOTYPE Erythromelana woodi Inclan D.J./ DI208MW. Additional specimens examined: Costa Rica, Bolivia, Mexico and Ecuador. [Examined by AJF.]
Described from 2 males and 2 females. Length: male = 6mm; female = 5–6mm.
Head: (Fig.
Neighbor-Joining (NJ –
General morphology of Erythromelana jimmychevezi sp. n. a–c: holotype male; voucher code: DHJPAR0038684; d–f: female paratype: DHJPAR0037257.
Thorax: (Fig.
Abdomen: dorsal surface of abdomen mostly black in ground color; ventral margins of T1+2 and T3 yellow in ground color, the yellow extending to 3/4 of lateral surface of these tergites; T4 yellow in ground color on 1/3–1/2 of lateral surface; in dorsal view, both T3 and T4 black in ground color medially and yellow laterally; ground color of T5 entirely black; transverse bands of sparse white pollinosisty present on anterior 1/3–1/4 of T3 and T4 and on anterior 2/3 of T5; one pair of median marginal setae on T1+2 and T3; T4 and T5 with a full row of marginal setae; mid-dorsal depression of T1+2 not reaching median marginal setae.
Male terminalia: (Fig.
This species is included in the E. cryptica species group (
Erythromelana jimmychevezi is named in recognition of Jimmy Chévez Elizondo for his contributions to the accounting team for Area de Conservación Guanacaste, the forest this fly lives in.
Costa Rica, ACG, Guanacaste Prov., rain forest, 470–675 m elevation.
A parasitoid of Eois Janzen52 (Geometridae), which feeds on Piper auritum (Piperaceae).
Described from 4 males and 4 females. Length: male = 6–7mm; female = 5–6mm .
Head: (Fig.
General morphology of Erythromelana glenriverai sp. n. a–c: holotype male; voucher code: DHJPAR0038680; d–f: female paratype; coucher code: DHJPAR0037426.
Thorax: (Fig.
Abdomen: ground color of dorsal surface mostly black; T1+2 all black in ground color in dorsal view, and of yellow ground color ventrally in lateral view; T3 mostly yellow on anterior 3/4, black posteriorly; coloration of T3, in dorsal view, appearing as a black triangle on a yellow background; T4 yellow in ground color on 1/3–1/2 of lateral surface, appearing as entirely black in dorsal view; T5 entirely black; transverse bands of sparse white pollinosity present on anterior 1/3–1/4 of T3 and T4, and on anterior 2/3 of T5; one pair of median marginal setae on T1+2 and T3; T4 and T5 with a full row of marginal setae; mid-dorsal depression of T1+2 not reaching median marginal setae.
Male terminalia: (Fig.
This species is included in the E. cryptica species group (
Erythromelana glenriverai is named in recognition of Glen Rivera Chaves for his contributions to the accounting team for Area de Conservación Guanacaste, the forest this fly lives in.
Costa Rica, ACG, Alajuela and Guanacaste Provs., rain forest, 430–740m elevation.
A parasitoid of Eois Janzen49, E. Janzen04, E. Janzen236 and E. dibapha (Schaus) (Geometridae), which feed on four species of rain forest Piperaceae.
The DNA barcode sequences recovered from the new Erythromelana species from ACG display the characteristic strong AT bias of insect mitochondrial DNA (mean percent GC content 32.89%, SE 0.08) and no insertions or deletions. Within-species variation was low (mean distance of 0.28%) compared to between-species variation (mean distance 5.18%). All values of DNA barcode variation were calculated within BOLD and can be re-calculated in the future as more specimens are recovered from the ACG inventory and added to the DNA library. Fig. 1 presents a neighbor–joining tree for the Erythromelana specimens reared and DNA barcoded by this inventory to date.
We gratefully acknowledge the unflagging support of the team of ACG parataxonomists (Janzen et al. 2009, Janzen & Hallwachs 2011) who collected 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, JRS Biodiversity Foundation, 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 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).