Biodiversity Data Journal :
Taxonomic Paper
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Corresponding author: AJ Fleming (ajfleming604@gmail.com)
Academic editor: Daniel Whitmore
Received: 16 Nov 2016 | Accepted: 25 Jan 2018 | Published: 02 Feb 2018
© 2018 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 AJ, Wood DM, Smith MA, Hallwachs W, Janzen DH (2018) Revision of the Mesoamerican species of Calolydella Townsend (Diptera: Tachinidae) and description of twenty-three new species reared from caterpillars in Area de Conservación Guanacaste, northwestern Costa Rica. Biodiversity Data Journal 6: e11223. https://doi.org/10.3897/BDJ.6.e11223
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Twenty-three new species of the genus Calolydella Townsend, 1927 (Diptera: Tachinidae) are described, all reared from multiple species of wild-caught caterpillars across a wide variety of families (Lepidoptera: Crambidae; Erebidae; Geometridae; Hesperiidae; Lycaenidae; Nymphalidae; Pieridae; Riodinidae; and Sphingidae). All caterpillars were collected within Area de Conservación Guanacaste (ACG), in northwestern Costa Rica. This study provides a concise description of each new species using morphology, life history, molecular data, and photographic documentation. In addition to the new species, we also provide a generic redescription and revised key to species of the genus Calolydella from Central and South America.
The following 23 new species of Calolydella are described by Fleming and Wood: C. adelinamoralesae sp. n., C. alexanderjamesi sp. n., C. argentea sp. n., C. aureofacies sp. n., C. bicolor sp. n., C. bifissus sp. n., C. crocata sp. n., C. destituta sp. n., C. discalis sp. n., C. erasmocoronadoi sp. n., C. felipechavarriai sp. n., C. fredriksjobergi sp. n., C. inflatipalpis sp. n., C. interrupta sp. n., C. nigripalpis sp. n., C. omissa sp. n., C. ordinalis sp. n., C. renemalaisei sp. n., C. susanaroibasae sp. n., C. tanyadapkeyae sp. n., C. tenebrosa sp. n., C. timjamesi sp. n., C. virginiajamesae sp. n. Lydella frugale Curran, 1934 is proposed as a new synonym of Pygophorinia peruviana Townsend, 1927, syn. n., under the combination Calolydella frugale (Curran, 1934), comb. n.
Blondeliini, caterpillars, cloud forest, dry forest, fly, Neotropics, parasitoid, rain forest
The New World genus Calolydella Townsend, 1927 (Tachinidae: Blondeliini) was erected for one male and several female specimens collected by Townsend himself at Itaquaquecetuba (now a suburb of São Paolo), Brazil and described as Calolydella geminata Townsend, 1927. When
Here, we describe 23 new species of Calolydella reared from wild-caught caterpillars collected as part of the ongoing inventory of Lepidoptera larvae (and their parasitoids) in the terrestrial portion of Area de Conservación Guanacaste (ACG), an area of 120,000 hectares in northwestern Costa Rica (
All flies and rearing information described herein were collected as part of the ongoing ACG inventory of caterpillars, their food plants, and their parasitoids, throughout the three major ACG terrestrial ecosystems (
Our present treatment of the genus Calolydella is limited to species from the Mesoamerican region, from the Mexican provinces of Guerrero and Oaxaca to the Colombian border with Panama. While we included all known species in our comparisons with the new species, only the species distributed within this region are included in our key.
AMNH American Museum of Natural History, New York, USA
NHMUK Natural History Museum [formerly British Museum (Natural History)], London, UK
CNC Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa, Canada
MNCR Museo Nacional de Costa Rica [formerly Instituto Nacional de Biodiversidad – INBio], Santo Domingo de Heredia, Costa Rica
NMW Naturhistorisches Museum Wien, Vienna, Austria
USNM National Museum of Natural History [formerly United States National Museum], Washington, D.C., USA
The management of voucher specimens has been detailed in previous papers in this series (
All DHJPARxxxxxxx-coded tachinids had one leg removed for DNA barcoding and couriered to the Center for Biodiversity Genomics of the Biodiversity Institute of Ontario (BIO) at the University of Guelph (Canada). All associated data and successful barcodes are permanently and publicly deposited in the Barcode of Life Data System (BOLD) (
All inventoried specimens discussed herein were collected under Costa Rican government research permits issued to DHJ; the Tachinidae samples were exported under permit by DHJ from Costa Rica to their final depository in the CNC. Tachinid identifications for the inventory are done by DHJ in coordination with a) visual inspection of morphology by AJF and DMW, b) DNA barcoding by MAS and BIO, and c) databasing and association with host caterpillars by DHJ and WH through the inventory itself.
The date of capture of fly specimens is the date of eclosion of the fly and not the date of capture of the caterpillar. The fly eclosion date is much more representative of the time when that species is on the wing than is the date of capture of the parasitized caterpillar. The “collector” is the parataxonomist who collected the caterpillar, rather than the person who later retrieved the newly eclosed fly and processed it by freezing, pinning, labeling and oven-drying. The biology and parasitization rates of individual tachinid species will be the subject of later papers. The holotypes of the new species described herein are deposited in the Diptera collection of the CNC, Ottawa, Canada.
Habitus and terminalia photographs were taken using the methods outlined in
Adult flies were dissected following standard practice (
Names of undescribed host species follow a standardized, interim naming system used for taxonomic units considered to be distinct species and identified by a combination of DNA barcode, morphology, and host-plant information. The interim names are given in the format "Eois Janzen52", where the species epithet is composed of the name of the taxonomist who first singled out the species and a number. This prevents confusion with already described species while maintaining traceability of each undescribed species within the ACG project.
We analyzed DNA barcodes (the 5’ region of the cytochrome c oxidase I (CO1) gene,
Calolydella Townsend, 1927a: 278. Type-species: Calolydella geminata Townsend, 1927, by original designation.
Olindopsis Townsend, 1927a: 274. Type-species: Olindopsis andinensis Townsend, 1927, by original designation. Synonymy proposed by Wood (1985).
Pygophorinia Townsend, 1927a: 274. Type-species: Pygophorinia peruviana Townsend, 1927, by original designation. Synonymy proposed by Wood (1985).
Prodexodes Townsend, 1927a: 280. Type-species: Prodexodes rufiventris Townsend, 1927, by original designation. Synonymy proposed by Wood (1985).
Calolydella belongs to the tribe Blondeliini.
The following redescription applies to both males and females of Calolydella; any differences between the sexes are noted. Head: males lacking proclinate orbital setae, females with 2 pairs of proclinate orbital setae; first reclinate orbital seta either equal to or longer than uppermost frontal seta; ocellar setae long; eye bare, if haired then hairs minute and inconspicuous; parafacial bare; facial margin level with vibrissa and not visible in profile, with a few small supra-vibrissal setae; subvibrissal ridge short, with three or fewer setae; margin of postgena straight or slightly convex anteriorly, genal dilation weak or absent; palps normally slender and yellow, some species with black or inflated palps; postpedicel of similar length in both sexes; arista minutely pubescent, slightly thickened at base. Thorax: prosternum setose; proepisternum bare; postpronotum with 2–4 setae in a narrow triangle or a straight line (rarely inner basal seta absent); acrostichal setae 1–3:3–4; dorsocentral setae 1–3:2–3; intra-alar setae 1–3:2–3; supra-alar setae 1–2:3; katepisternal setae 2–3 [
Other species included in Calolydella Townsend
andinensis Townsend, 1927: 339 (Olindopsis). Holotype female (USNM), by original designation [examined by DMW]. Type locality: Peru, Cordillera Oriental, near Tabalosas, 3000ft.
blandita Wulp, 1890: 142 (Hypostena). Lectotype male (NHMUK), by fixation of Wood (1985:28). Type locality: Mexico, Guerrero, Sierra de las Aguas Escondidas, 7000ft.
cingulata Schiner, 1868: 327 (Meigenia). Holotype male (NMW), by original designation [examined by DMW]. Type locality: Brazil.
cylindriventris Wulp, 1890: 145 (Hypostena). Lectotype male (NHMUK), by fixation of Wood (1985:28). Type locality: Mexico, Guerrero, Sierra de las Aguas Escondidas, 7000ft.
geminata Townsend, 1927: 293 (Calolydella). Multiple syntypes (USNM) [1 female and several males examined by DMW]. Type locality: Brazil, São Paulo, Itaquaquecetuba.
gentica Walker, 1860: 302 (Masicera). Lectotype male (NHMUK), by fixation of Wood (1985:28) (examination of “Holotype: ♂” from Mexico in NHMUK is regarded as a lectotype fixation [JE O'Hara, pers. comm.]) [examined by DMW]. Type locality: Mexico.
lathami Curran, 1925: 284 (Lydella). Holotype male (CNC), by original designation [examined by DMW]. Type locality: USA, New York, Long Island, Greenpoint.
leucophaea Wulp, 1890: 141 (Hypostena). Lectotype female (NHMUK), by fixation of Wood (1985:28). Type locality: Mexico, Guerrero, Sierra de las Aguas Escondidas, 9500ft.
peruviana Townsend, 1927: 355 (Pygophorinia). Holotype female (USNM), by original designation [examined by DMW]. Type locality: Peru, [Puno], Casahuiri.
frugale Curran, 1934: 511 (Lydella). Holotype female (AMNH), by original designation [examined by DMW]. Type locality: Guyana, Bartica, Kartabo. Syn n., comb. n.
rufiventris Townsend, 1927: 350 (Prodexodes). Holotype male (USNM), by original designation [examined by DMW]. Type locality: Brazil, Sao Paulo, Itaquaquecetuba.
summatis Reinhard, 1974: 1158 (Calolydella). Holotype male (CNC), by original designation [examined by DMW]. Type locality: Mexico, Durango, 14 miles southwest of El Salto, 8000ft.
triangulifera Bigot, 1889: 268 (Homodexia). Holotype male (NHMUK), by monotypy [examined by DMW]. Type locality: Mexico.
pictigaster Bigot, 1889: 261 (Ceromasia). Holotype female (NHMUK), by monotypy [examined by DMW]. Type locality: Mexico (NHUMK).
trifasciata Walker, 1837: 350 (Tachina). Lectotype female (NHMUK), by fixation of Wood (1985:29) (examination of “Holotype: ♀” from South America in NHMUK is regarded as a lectotype fixation [JE O'Hara, pers. comm.]) [examined by DMW]. Type locality: South America, precise locality uncertain (
quadrivittata Bigot, 1889: 261 (Ceromasia). Holotype male (NHMUK) [published as female], by monotypy [examined by DMW]. Type locality: Mexico.
quadristriata Wulp, 1890: 146 (Hypostena). Holotype male (NHMUK), by original designation [examined by DMW]. Type locality: Costa Rica, Volcán de Irazu, 6000ft.
Calolydella are slender-bodied flies characterized by an abdomen longer than it is wide, with bold abdominal markings of yellow-gold pollinose stripes on a black background. According to
Essentially a Neotropical genus save for one species, Calolydella lathami, which is present in the eastern USA and adjacent Canada.
All species of Calolydella are parasitoids of Lepidoptera larvae in the families Crambidae, Erebidae, Geometridae, Hesperiidae, Lycaenidae, Notodontidae, Nymphalidae, Pieridae, Riodinidae, and Sphingidae.
Male (Fig.
General morphology of Calolydella adelinamoralesae sp. n.; a–c: male holotype, voucher n. DHJPAR0017775; d–f: female paratype, voucher n. DHJPAR0038669.
Female (Fig.
Calolydella adelinamoralesae can be distinguished from all other species of Calolydella by the following combination of traits: parafacial all silver, frontal setae extending to base of postpedicel, anatergite bare, and T4 with 1–2 pairs of discal setae.
The specific epithet is in honor of Adelina Morales Chaves of La Cruz, Guanacaste, Costa Rica, in recognition of the many years she dedicated to sorting ACG Malaise trap samples, many of which contained Calolydella flies.
Costa Rica, ACG, Alajuela and Guanacaste provinces, 96–527m.
Calolydella adelinamoralesae has been reared 53 times from Agaraea minuta (Schaus, 1892) (Lepidoptera: Erebidae), in rain forest and dry-rain lowland intergrade.
Male (Fig.
Female: not known at this time.
Calolydella alexanderjamesi can be distinguished from all other species of Calolydella by the following combination of traits: parafacial mostly gold, fronto-orbital plate with small black setulae interspersed among frontal setae, anatergite bare, and thoracic pollinosity gold on both dorsal and lateral surfaces.
The specific epithet is in honor of Alexander James of Levittown, Pennsylvania, in recognition of the moral and family support of his mother, Tanya Dapkey, in her efforts curating and preparing ACG parasitoid flies for DNA barcoding.
Costa Rica, ACG, Guanacaste Province, Area Administrativa, 295m.
Calolydella alexanderjamesi has been reared once from Correbia undulata (Druce, 1884) (Lepidoptera: Erebidae), in dry forest.
Female (Fig.
Male: not known at this time.
Calolydella argentea can be distinguished from all other species of Calolydella by the following combination of traits: parafacial and fronto-orbital plate entirely silver, frontal setae extending to base of postpedicel with intermingled small setulae, 3:3 acrostichal setae, anterior 2/3 of abdominal tergites with silver pollinose bands, anatergite with three or more hair-like setae, and wing vein R4+5 with 6–7 setulae, almost reaching to crossvein R-M.
The specific epithet is derived from the Latin adjective "argentum", meaning silver, in reference to its entirely silver parafacial and fronto-orbital plate, a feature unique to this species.
Costa Rica, ACG, Alajuela Province, Finca Esmeralda, 123m.
Calolydella argentea has been reared once from Antiblemma Poole33 (Lepidoptera: Noctuidae), in rain forest.
Female: 6mm (Fig.
Male: not known at this time.
Calolydella aureofacies can be distinguished from all other species of Calolydella by the following combination of traits: parafacial mostly gold, female palps black basally, anatergite with three or more hair-like setae, and scutellar discal setae absent.
The specific epithet is derived from the Latin adjective "aurum", meaning gold, and "facies", meaning face, in reference to its gold parafacial and fronto-orbital plate.
Costa Rica, ACG, Guanacaste Province, Sendero Ponderosa, 1060m.
Calolydella aureofacies has been reared once from Cyclophora Janzen23 (Lepidoptera: Geometridae), in cloud forest.
Female (Fig.
Male: not known at this time.
Calolydella bicolor can be distinguished from all other species of Calolydella by the following combination of traits: fronto-orbital plate entirely gold, parafacial entirely silver, anatergite with three or more hair-like setae, and gold abdominal pollinose bands interrupted by a median dark stripe.
The specific epithet is derived from the Latin adjective “bicolor”, meaning two-toned, in reference to its gold fronto-orbital plate and silver parafacial.
Costa Rica, ACG, Guanacaste Province, Sendero Bernales, 660m.
Calolydella bicolor has been reared once from Callicore lyca (Doubleday, 1847) (Lepidoptera: Nymphalidae), in rain forest.
Male (Fig.
General morphology of Calolydella bifissus sp. n.; a–c: male holotype, voucher n. DHJPAR0017132; d–f: female paratype, voucher n. 07-SRNP-40315.
Female (Fig.
Calolydella bifissus can be distinguished from all other species of Calolydella by the following combination of characters: two bold thoracic vittae, abdominal pollinosity concolorous on both dorsal and ventral surfaces, anatergite with a small tuft of hair-like setae, and abdominal ground color dark brown-orange.
The specific epithet is derived from the Latin adjective "bifissus", meaning cleft or cloven, in reference to cleft nature of the cerci, which are fused basally, a character state unique to this species.
Costa Rica, ACG, Alajuela and Guanacaste provinces, 400–1150m.
Calolydella bifissus has been reared twice, from two separate hosts, Dircenna kluggii (Geyer, 1837) (Lepidoptera: Nymphalidae) and Phyllodonta latrata (Guenée, 1857) (Lepidoptera: Geometridae), in rain forest and cloud forest ecosystems.
Male (Fig.
Female: not known at this time.
Calolydella crocata can be distinguished from all other species of Calolydella by the following combination of traits: fronto-orbital plate mostly gold-pollinose but with some silver pollinosity and sparsely setulose, thorax with four vittae visible only under certain angles of light, anatergite bare, and abdomen with transverse marginal pollinose bands interrupted by a thin dark median stripe.
The specific epithet is derived from the Latin adjective "crocatum", meaning yellow, in reference to the gold pollinosity present over most of the body of the new species.
Costa Rica, ACG, Guanacaste Province, Estación Quica, 470m.
Calolydella crocata has been reared once from Semaeopus Janzen05 (Lepidoptera: Geometridae), in rain forest.
Male (Fig.
General morphology of Calolydella destituta sp. n.; a–c: male holotype, voucher n. DHJPAR0017799; d–f: female paratype, voucher n. DHJPAR0017771.
Female (Fig.
Calolydella destituta can be distinguished from all other species of Calolydella by the following combination of traits: anatergite with a small tuft of setae, pollinose bands on abdomen not interrupted by a dark median stripe, and T3 lacking median marginal setae.
The specific epithet is derived from the Latin adjective "destituta", meaning absent or lacking, in reference to the absence of median marginal setae on T3, a character state unique to this species.
Costa Rica, ACG, Guanacaste Province, 345.
Calolydella destituta has been reared four times from Ramphia albizona (Latreille, 1817) (Lepidoptera: Erebidae), in rain forest and dry-rain lowland intergrade.
Male (Fig.
Female: not known at this time.
Calolydella discalis can be distinguished from all other species of Calolydella by the following combination of traits: parafacial mostly gold, fronto-orbital plate with a single row of small black setulae outside of the frontal setae, anatergite with a small tuft of three or more hair-like setae, and T3 and T4 each with two pairs of discal setae.
The specific epithet refers to the two pairs of discal setae on T3 and T4.
Costa Rica, ACG, Alajuela Province, Rio Blanco Abajo, 500m.
Calolydella discalis has been reared twice from Turuptiana obliqua Walker, 1869 (Lepidoptera: Erebidae: Arctiinae), in rain forest.
Male (Fig.
General morphology of Calolydella erasmocoronadoi sp. n.; a–c: male holotype, voucher n. DHJPAR0039246; d–f: female paratype, voucher n. 04-SRNP-55202.
Female (Fig.
Calolydella erasmocoronadoi can be distinguished from all other species of Calolydella by the following combination of traits: fronto-orbital plate bare, up to 50% silver pollinose, thoracic vittae visible only under certain angles of light, anatergite with three or more setae arranged in a small tuft, wing vein R4+5 with 6–7 setulae, and abdomen with uninterrupted transverse marginal pollinose bands.
The specific epithet is in honor of Erasmo Coronado Caballo of Liberia, Costa Rica, in recognition of his high-quality assistantship to Sigifredo Marin of Liberia and to the ACG parataxonomists who collected and reared the caterpillars from which the new species of ACG Calolydella described here emerged.
Costa Rica, ACG, Guanacaste Province, Sendero Mismo, 680m.
Calolydella erasmocoronadoi has been reared 42 times from Euselasia chryssipe (Bates, 1866) (Lepidoptera: Riodinidae), in rain forest.
Male (Fig.
General morphology of Calolydella felipechavarriai sp. n.; a–c: male holotype, voucher n. DHJPAR0050491; d–f: female paratype, voucher n. DHJPAR0050489.
Female (Fig.
Calolydella felipechavarriai can be distinguished from all other species of Calolydella by the following combination of traits: fronto-orbital plate almost bare, anatergite with at most two hair-like setae, and T4 with one pair of discal setae.
The specific epithet is in honor of Luis Felipe Chavarria Diaz of ACG, Guanacaste, Costa Rica, in recognition of his high quality accounting, photography and administrative support of the Guanacaste Dry Forest Conservation Fund (www.gdfcf.org) and of the ACG parataxonomists who collected and reared the caterpillars from which the new species of ACG Calolydella described here emerged.
Costa Rica, ACG, Alajuela Province, Palomo, 96m.
Calolydella felipechavarriai has been reared twice from Bardaxima perses Druce, 1900 (Lepidoptera: Notodontidae), in rain forest.
Male (Fig.
General morphology of Calolydella fredriksjobergi sp. n.; a–c: male holotype, voucher n. DHJPAR0011711; d–f: female paratype, voucher n. DHJPAR0054013.
Female (Fig.
Calolydella fredriksjobergi can be distinguished from all other species of Calolydella by the following combination of characters: parafacial mostly gold, anatergite with three or more hair-like setae arranged in a small tuft, two pairs of discal setae on T3 and one pair of discal setae on T4.
The specific epithet is in honor of Fredrik Sjoberg of Runmaro Island, Sweden, in recognition of his wonderful book "The Fly Trap" (
Costa Rica, ACG, Alajuela and Guanacaste provinces, 105–1060m.
Calolydella fredriksjobergi has been reared six times from six different species belonging to various families of Lepidoptera (Nymphalidae; Erebidae; and Crambidae), in rain forest, cloud forest, and dry forest ecosystems.
Male (Fig.
General morphology of Calolydella inflatipalpis sp. n.; a–c: male holotype, voucher n. DHJPAR0016181; d–f: female paratype, voucher n. DHJPAR0016169.
Female (Fig.
Calolydella inflatipalpis can be distinguished from all other species of Calolydella by the following combination of traits: fine setulae interspersed among frontal setae, palps normal in male but significantly inflated in female, anatergite with three or more hair-like setae arranged in a small tuft, and T3 and T4 each with two pairs of discal setae.
The specific epithet is derived from the Latin adjective “inflatum”, meaning inflated, and "palpus", meaning palp, in reference to the inflated, bulbous palpi of females of this species.
Costa Rica, ACG, Alajuela and Guanacaste provinces, 290–1150m.
Calolydella inflatipalpis has been reared 31 times from four separate species of Heliconiini: Dione juno (Cramer, 1779), Dryas iulia (Fab., 1775), Dryadula phaetusa (Linn., 1758), and Heliconius charithonia (Linn., 1767) (Lepidoptera: Nymphalidae), in rain forest, dry forest, cloud forest and dry-rain lowland intergrade ecosystems.
Male (Fig.
General morphology of Calolydella interrupta sp. n.; a–c: male holotype, voucher n. DHJPAR0029598; d–f: female paratype, voucher n. 08-SRNP-36564.
Female (Fig.
Calolydella interrupta can be distinguished from all other species of Calolydella by the following combination of traits: parafacial at least 50% silver pollinose, two bold thoracic vittae, abdominal pollinosity interrupted by median stripe, anatergite with three or more setae arranged in a small tuft, and T3 with a complete row of marginal setae.
The specific epithet is derived from the Latin adjective “interruptum", meaning breach or interruption, in reference to dark median stripe breaking up the gold marginal pollinosity on the abdomen of this species.
Costa Rica, ACG, Guanacaste Province, Sendero Arenales 1080m.
Calolydella interrupta has been reared once from Acharia ophelians (Dyar, 1927) (Lepidoptera: Limacodidae), in cloud forest.
Male (Fig.
Female: not known at this time.
Calolydella nigripalpis can be distinguished from all other species of Calolydella by the following combination of characters: frontal setae not extending beyond base of postpedicel, palps black in male, four barely visible thoracic vittae, anatergite bare, and abdominal ground color black.
The specific epithet is derived from the Latin adjective "nigrum", meaning black, and "palpus", meaning palp, in reference to the black palpi of males of this species.
Costa Rica, ACG, Alajuela and Guanacaste provinces, 135–520m.
Calolydella nigripalpis has been reared six times from Semaeopus Janzen08 (Lepidoptera: Geometridae), in rain forest.
Male (Fig.
General morphology of Calolydella omissa sp. n.; a–c: male holotype, voucher n. DHJPAR0016605; d–f: female paratype, voucher n. 06-SRNP-43825.
Female (Fig.
Calolydella omissa can be distinguished from all other species of Calolydella by the following combination of traits: fronto-orbital plate gold, parafacial silver, abdomen with uninterrupted transverse silver pollinose bands across all tergites, anatergite bare (rarely setose), and T3 without discal setae.
The specific epithet is derived from the Latin adjective "omissa", meaning dropped or absent, in reference to the absence of discal setae on T3, a character state unique to this species.
Costa Rica, ACG, Alajuela Province, Cabanya, 340m.
Calolydella omissa has been reared once from Letis Janzen03 (Lepidoptera: Erebidae), in rain forest.
Male (Fig.
General morphology of Calolydella ordinalis sp. n.; a–c: male holotype, voucher n. DHJPAR0017787; d–f: female paratype, voucher n. DHJPAR0017792.
Female (Fig.
Calolydella ordinalis can be distinguished from all other species of Calolydella by the following combination of characters: parafacial mostly gold, fronto-orbital plate with setulae sparsely interspersed among frontal setae, anatergite bare, and a complete row of discal setae on T4.
The specific epithet is derived from the Latin adjective "ordinarius", meaning ordinary or complete, in reference to the complete, regular row of discal setae on T4, a character state unique to this species.
Costa Rica, ACG, Guanacaste Province, Sendero Maritza, 760m.
Calolydella ordinalis has been reared six times from Eloria torrida Schaus, 1910 (Lepidoptera: Erebidae) in cloud forest.
Male (Fig.
Female: not known at this time.
Calolydella renemalaisei can be distinguished from all other other species of Calolydella by the following combination of characters: frontal setae extending to base of postpedicel, fronto-orbital plate sparsely setose througout, with four thoracic vittae, the outermost two twice as wide as the innermost two, anatergite with three or more hair-like setae arranged in a small tuft, and up to two pairs of discal setae on T4.
The specific epithet is in honor of René Edmond Malaise (1892-1978) of Stockholm, Sweden, in recognition of his invention of the Malaise trap, which has captured so many ACG Tachinidae and other insects, as explained by Fredrik Sjöberg in his wonderful book "The Fly Trap" (
Costa Rica, ACG, Guanacaste Province, 280–680m.
Calolydella renemalaisei has been reared six times from two species of Arctiinae: Euceron aeolum Hampson, 1898, and Leucotmemis nexa Herrich-Schäffer, 1854 (Lepidoptera: Erebidae), in rain forest and dry-rain lowland intergrade ecosystems.
Male (Fig.
General morphology of Calolydella susanaroibasae sp. n.; a–c: male holotype, voucher n. DHJPAR0057699; d–f: female paratype, voucher n. 15-SRNP-1269.
Female (Fig.
Calolydella susanaroibasae can be distinguished from all other species of Calolydella by the following combination of characters: fronto-orbital plate with a single row of fine setulae outside of frontal setae, thoracic pollinosity gold dorsally and on 50% of lateral surface, with outer two thoracic vittae twice as wide as inner two, anatergite with three or more hair-like setae arranged in a small tuft, and base of ST1+2 black lateroventrally.
The specific epithet is in honor of Susana Roibas of Lexington, Kentucky, in recognition of her company "Sante-Traps" (http://www.santetraps.com), which has long supplied the world with Malaise traps and other kinds of insect traps, many of which have been used to collect ACG Tachinidae and other insects.
Costa Rica, ACG, Alajuela Province, Rio Blanco Abajo, 500m.
Calolydella susanaroibasae has been reared once from Isanthrene echemon Druce, 1884 (Lepidoptera: Erebidae: Arctiinae), in rain forest.
Female (Fig.
Male: not known at this time.
Calolydella tanyadapkeyae can be distinguished from all other other species of Calolydella by the following combination of traits: parafacial all silver, frontal setae extending to base of postpedicel, thoracic pollinosity gold dorsally and over 50% of lateral surfaces, postpronotum with only two setae (inner basal seta absent), anatergite bare, and scutellar discal setae situated as wide apart as subapical scutellar setae.
The specific epithet is in honor of Tanya Dapkey James of Levittown, Pennsylvania, in recognition of her efforts curating and preparing ACG parasitoid flies for DNA barcoding.
Costa Rica, ACG, Alajuela Province, Camino Rio Francia, 410m.
Calolydella tanyadapkeyae has been reared once from Uranophora flaviceps (Hampson, 1901) (Lepidoptera: Erebidae), in rain forest.
Male (Fig.
General morphology of Calolydella tenebrosa sp. n.; a–c: male holotype, voucher n. DHJPAR0017810; d–f: female paratype, voucher n. DHJPAR0053996
Female (Fig.
Calolydella tenebrosa can be distinguished from all other species of Calolydella by the following combination of characters: frontal setae extending beyond base of postpedicel, palps black in males and dark grayish-orange in females, abdominal ground color dark brown-orange, anatergite bare, and abdominal T5 black apically in both sexes.
The specific epithet Calolydella tenebrosa is derived from the Latin adjective "tenebrosum", meaning black or dark, in reference to the dark-colored palpi and dark-colored legs, character states found in both sexes of this species.
Costa Rica, ACG, Alajuela and Guanacaste provinces, 320–380m.
Calolydella tenebrosa has been reared once from Cyclophora Janzen14 and once from Semaeopus Janzen08 (Lepidoptera: Geometridae), in rain forest.
Male (Fig.
General morphology of Calolydella timjamesi sp. n.; a–c: male holotype, voucher n. DHJPAR0017781; d–f: female paratype, voucher n. DHJPAR0016192.
Female (Fig.
Calolydella timjamesi can be distinguished from all other species of Calolydella by the following combination of traits: fronto-orbital plate sparsely setulose to bare, frontal setae extending beyond base of postpedicel, abdominal ground color black, anatergite with three or more setae often in a small tuft, and scutellar discal setae slightly closer togetherthan subapical scutellar setae.
The specific epithet is in honor of Tim James of Levittown, Pennsylvania, in recognition of the moral and family support of Tanya Dapkey in her efforts curating and preparing ACG parasitoid flies for DNA barcoding.
Costa Rica, ACG, Alajuela and Guanacaste provinces, 335–740m.
Calolydella timjamesi has been reared 26 times from six species of Notodontidae: Dunama janewaldronae Chacon, 2013, Dunama jessiehillae Chacon, 2013, Tithraustes noctilucesICG02, Tithraustes lambertae Miller, 2008, Dioptis longipennis (Schaus, 1913), and Dottia viridifusca (Schaus, 1906), in rain forest and dry-rain lowland intergrade.
Male (Fig.
General morphology of Calolydella virginiajamesae sp. n.; a–c: male holotype, voucher n. DHJPAR0017770; d–f: female paratype, voucher n. DHJPAR0045648.
Female (Fig.
Calolydella virginiajamesae can be distinguished from all other species of Calolydella by the following combination of traits: fronto-orbital plate with a single row of small black setulae outside of frontal setae, parafacial silver-pollinose, anatergite with three or more setae arranged in a small tuft, and T3–T4 each with a single pair of discal setae.
The specific epithet is in honor of Virginia James of Levittown, Pennsylvania, in recognition of the moral and family support of her mother, Tanya Dapkey, in her efforts curating and preparing ACG parasitoid flies for DNA barcoding.
Costa Rica, ACG, Alajuela and Guanacaste provinces, 85–1150m.
Calolydella virginiajamesae has been reared 34 times from various species of Lepidoptera from several families (Nymphalidae; Hesperiidae; Lycaenidae; Geometridae; Crambidae; Erebidae; Riodinidae; Sphingidae; and Pieridae), in rain forest, cloud forest, dry forest, and dry-rain lowland intergrade ecosystems.
Key to the species of Calolydella of the Mesoamerican region This key is confined to species of Calolydella occurring in the Mesoamerican biogeographical region. It is applicable to both males and females. |
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1 | Fronto-orbital plate with gold pollinosity; thorax with some gold on both lateral and dorsal surfaces | 2 |
– | Fronto-orbital plate entirely silver; thorax silver on both lateral and dorsal surfaces | 25 |
2 | Fronto-orbital plate entirely gold | 3 |
– | Fronto-orbital plate with both silver and gold pollinosity (Fig. |
Calolydella erasmocoronadoi sp. n. |
3 | Abdomen with transverse marginal pollinose bands interrupted by a median dark stripe | 4 |
– | Abdomen with uninterrupted transverse marginal pollinose bands | 8 |
4 | Parafacial pollinosity either entirely silver or up to 50% silver pollinose | 5 |
– | Parafacial pollinosity entirely gold | Calolydella gentica (Walker, 1860) |
5 | Fronto-orbital plate with setulae outside of frontal seta; thorax with four distinct dorsal vittae; anatergite with up to two fine hair-like seta; T3 with one pair of median marginal setae | 6 |
– | Fronto-orbital plate bare; thorax with two, prominent, dark dorsal vittae; anatergite with three or more hair-like setae; T3 with a complete row of marginal setae and one pair of discal setae | 7 |
6 | Fronto-orbital plate sparsely setulose; frontal setae not extending beyond base of postpedicel; anatergite bare; wing vein R4+5 with 4–5 small setulae dorsally, near base; T3 with one pair of discal setae (Fig. |
Calolydella crocata sp. n. |
– | Fronto-orbital plate with a row of fine setulae outside of frontal setae; frontal setae extending beyond base of postpedicel; anatergite with up to two hair-like setae; wing vein R4+5 with at most three small setulae dorsally, near base; T3 with two pairs of discal setae | Calolydella trifasciata (Walker, 1837) |
7 | Thorax with two presutural acrostichal setae; abdominal ST1+2 mostly black dorsally, with small spots of gold pollinosity laterally (Fig. |
Calolydella interrupta sp. n. |
– | Thorax with three presutural acrostichal setae; abdominal ST1+2 with a thick median black stripe but still mostly gold dorsally (Fig. |
Calolydella bicolor sp. n. |
8 | T3 with at least one pair of median marginal setae | 9 |
– | T3 without median marginal setae (Fig. |
Calolydella destituta sp. n. |
9 | T3 with at least one pair of discal setae | 10 |
– | T3 without discal setae (Fig. |
Calolydella omissa sp. n. |
10 | Palpus dark gray to black | 11 |
– | Palpus yellow-orange | 12 |
11 | Frontal setae extending to base of postpedicel but not beyond; thoracic vittae only faintly visible; abdominal ground color black (Fig. |
Calolydella nigripalpis sp. n. |
– | Frontal setae extending beyond base of postpedicel; thoracic vittae prominent and easily visible; abdominal ground color dark brown-orange (Fig. |
Calolydella tenebrosa sp. n. |
12 | One or more pairs of discal setae on abdominal T4 | 13 |
– | A complete row of discal setae on abdominal T4 (Fig. |
Calolydella ordinalis sp. n. |
13 | Frontal setae extending beyond base of postpedicel | 14 |
– | Frontal setae extending to base of postpedicel | 17 |
14 | Parafacial silver; palpus normal; T3 and T4 each with one pair of discal setae | 15 |
– | Parafacial gold; palpus normal in male but significantly enlarged in female (Fig. |
Calolydella inflatipalpis sp. n. |
15 | Four distinct thoracic vittae postsuturally | 16 |
– | Thoracic vittae fused postsuturally (Fig. |
Calolydella timjamesi sp. n. |
16 | Anatergite with up to two hair-like setae or, rarely, bare (Fig. |
Calolydella felipechavarriai sp. n. |
– | Anatergite with a small tuft of three or more hair-like setae (Fig. |
Calolydella virginiajamesae sp. n. |
17 | Abdominal pollinosity concolorous on both ventral and dorsal surfaces | 18 |
– | Abdominal pollinosity gold dorsally, silver ventrally | 19 |
18 | Parafacial all gold pollinose; palpus black basally; thorax with four regular thoracic vittae; scutellum without discal setae; abdomen silver pollinose on both dorsal and ventral surfaces (Fig. |
Calolydella aureofacies sp. n. |
– | Parafacial with some silver pollinosity; palpus orange; thorax with two bold and prominent thoracic vittae; scutellum with discal setae; abdomen gold pollinose on both dorsal and ventral surfaces (Fig. |
Calolydella bifissus sp. n. |
19 | Base of ST1+2 with an orange spot lateroventrally | 20 |
– | Base of ST1+2 black lateroventrally (Fig. |
Calolydella tanyadapkeyae sp. n. |
20 | Parafacial silver-pollinose throughout | 21 |
– | Parafacial with both silver and gold pollinosity | 22 |
21 | Anatergite with three or more hair-like setae arranged in a tuft; thoracic chaetotaxy as follows: 3:3 acrostichal setae; 3:3 dorsocentral setae; 2:3 intra-alar setae; 2:3 supra-alar setae; abdominal T4 with one pair of discal setae (Fig. |
Calolydella fredriksjobergi sp. n. |
– | Anatergite bare; thoracic chaetotaxy as follows: 3:3 acrostichal setae (2:3 in females); 3:4 dorsocentral setae; 1:3 intra-alar setae; 1:3 supra-alar setae; abdominal T4 with two pairs of discal setae(Fig. |
Calolydella adelinamoralesae sp. n. |
22 | Outermost thoracic vittae 2X as wide as innermost, these becoming fused postsuturally | 23 |
– | Four thoracic vittae of sub-equal width, not fused postsuturally | 24 |
23 | Fronto-orbital plate with a single row of setulae outside of frontal setae; postpronotum with three setae; wing vein R4+5 with 5–7 setulae dorsally, reaching from base almost to crossvein R-M; abdominal T4 with two pairs of discal setae (Fig. |
Calolydella susanaroibasae sp. n. |
– | Fronto-orbital plate with setulae interspersed among frontal setae; postpronotum with two setae, inner basal seta absent; wing vein R4+5 with at most three small setulae dorsally at base; T4 with one pair of discal setae (Fig. |
Calolydella alexanderjamesi sp. n. |
24 | Fronto-orbital plate with a single row of setulae outside of frontal setae; thoracic chaetotaxy as follows: 3:3 acrostichal setae; 2:3 dorsocentral setae; 3:3 intra-alar setae; 2:3 supra-alar setae; wing vein R4+5 with more than 4–5 setulae dorsally, which may or may not extend to crossvein R-M; T3 and T4 both with up to two pairs of discal setae (Fig. |
Calolydella discalis sp. n. |
– | Fronto-orbital plate with setulae interspersed among frontal setae; thoracic chaetotaxy as follows: 3:3 acrostichal setae; 3:3 dorsocentral setae; 1:3 intra-alar setae; 2:3 supra-alar setae; wing vein R4+5 with at most 2–3 setulae dorsally at base, never extending to crossvein R-M; T3 with 1 pair of discal setae; T4 with up to two pairs of discal setae (Fig. |
Calolydella renemalaisei sp. n. |
25 | Thorax with two longitudinal vittae | 26 |
– | Thorax with four distinct longitudinal vittae | 28 |
26 | T3 with only one pair of discal setae; abdominal pollinosity appearing as thick uninterrupted bands across tergites | 27 |
– | T3 with two pairs of discal setae; abdominal pollinosity across tergites interrupted by a dark median stripe | Calolydella summatis Reinhard, 1974 |
27 | Frontal setae extending to base of postpedicel; three supravibrissal setae; 3:3 acrostichal setae; anatergite with three or more hair-like setae in a small tuft; anterior 2/3 of abdominal tergites silver pollinose; wing vein R4+5 with 6–7 setulae dorsally, extending almost to crossvein R-M (Fig. |
Calolydella argentea sp. n. |
– | Frontal setae extending beyond base of postpedicel; six or more supravibrissal setae; 2:3 acrostichal setae; anatergite bare; anterior 1/3 of abdominal tergites silver pollinose; wing vein R4+5 with two setulae dorsally, not extending to crossvein R-M | Calolydella cylindriventris (Wulp, 1890) |
28 | Fronto-orbital plate bare; frontal setae reaching base of postpedicel; thoracic pollinosity silver throughout; T3 with one pair of discal setae and one pair of median marginal setae | Calolydella triangulifera (Bigot, 1889) |
– | Fronto-orbital plate with sparse setulae interspersed among frontal setae; frontal setae reaching beyond base of postpedicel; thoracic pollinosity gold throughout; T3 with two pairs of discal setae and a row of marginal setae | Calolydella blandita (Wulp, 1890) |
A phylogenetic tree based on DNA barcodes was used to visually demonstrate the variation within and between species, and is presented in Fig.
Maximum Likelihood tree showing DNA barcode variation within ACG species of Calolydella. The inter- and intra-specific variation within the DNA barcode region for the 23 species of Calolydella from ACG is illustrated using a Maximum Likelihood tree based on the General Time Reversible model (GTR) (
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 are home to 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 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).