Zootaxa 3737 (1): 068–076 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2013 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3737.1.5 http://zoobank.org/urn:lsid:zoobank.org:pub:24550C94-58BD-493A-8BB3-3F7F64CDF703 A new species of Lixophaga Townsend (Diptera: ) from Colombia, a parasitoid of Neoleucinodes elegantalis (Guenée) (Lepidoptera: Crambidae)

NANCY CARREJO1, ANA E. DIAZ2 & NORMAN E. WOODLEY3,4 1Universidad del Valle, Departamento de Biología, A.A. 25630, Cali, Colombia. E-mail: [email protected] 2CORPOICA, La Selva, Rionegro Antioquia, Colombia. E-mail: [email protected] or [email protected] 3Systematic Entomology Laboratory, ARS, USDA, c/o Smithsonian Institution NHB-168, P. O. Box 37012, Washington DC 20013- 7012, USA. E-mail: [email protected] 4Corresponding author

Abstract

A new species of Lixophaga Townsend (Diptera: Tachinidae) from Colombia, Lixophaga puscolulo Carrejo & Woodley, sp. nov., is described and illustrated. It is a parasitoid of the tomato fruit borer, Neoleucinodes elegantalis (Guenée) (Lep- idoptera: Crambidae), an pest of Solanum quitoense Lam., in Colombia. Aspects of its biology are briefly discussed.

Key words: Diptera, Tachinidae, Lixophaga puscolulo, new species, Neotropical Region

Introduction

In the New World, the genus Lixophaga Townsend is one of the most species-rich genera of (Wood 1985). The genus has a wide distribution range which extends from much of the New World to the Oriental and Palearctic regions (Byun & Han 2011). In the New World, there are currently 64 described species of Lixophaga (Guimarães 1977; Wood 1985). Prior to now, no species of Lixophaga has been recorded from Colombia, but numerous species must occur there. There is very little taxonomic information on Neotropical Lixophaga other than the summary provided by Wood (1985), making it a difficult group to study. Arnaud (1978), Crosskey (1973), Guimarães (1977) and Shima (2006) list 28 genera of Lepidoptera and 10 genera of Coleoptera that have been reported as insect hosts of about 14 species of Lixophaga. Recently, Byun & Han (2011) presented a diagnosis of Lixophaga listing a number of character states. However, these character states individually are found in other Blondeliini, so only the combination of these character states can be considered diagnostic for the genus. Wood (1985) provided a more detailed diagnosis of Lixophaga, and mentioned the presence of a large seta on each side of sternite 5 in the male of at least some Lixophaga species, which is probably apomorphic for that group of species. The species of Lixophaga described below possesses these setae. For practical purposes, the keys to genera in Wood (1985) and Wood & Zumbado (2010) can be used to identify Lixophaga. The tomato fruit borer, Neoleucinodes elegantalis (Guenée) (Lepidoptera: Crambidae), is a Neotropical oligophagous pest which is mainly associated with cultivated species of the family Solanaceae, such as Capsicum annuum L. (green and red pepper), Solanum quitoense Lam. (lulo or narajanjilla), S. betaceum Cav. (tree tomato), S. lycopersicum Lam. (tomato), S. melongena L. (eggplant), S. sessiliflorum Dunal (cocona), S. pseudolulo Heiser (pacific lulo) and other wild species of Solanaceae (Capps 1948; A.L.A.E 1968; Sánchez 1973; Posada et al. 1981; Gallego & Velez 1992; Viáfara et al. 1999; Morales et al. 2003; Diaz 2009, 2010; Revelo et al. 2010; Anteparra et al. 2010). Here we describe a new species of Lixophaga, L. puscolulo Carrejo & Woodley, sp. nov., a parasitoid which was reared from the last instar larvae of N. elegantalis. The parasitoid has been reared only from the host associated with S. quitoense and from the wild solanaceous plant S. hirtum Vahl in Colombia (Diaz & Brochero 2012).

68 Accepted by J. O'Hara: 4 Nov. 2013; published: 18 Nov. 2013 The females of L. puscolulo that we observed in the field apparently are guided by odors of the volatile substances present in the excrement of the larvae of N. elegantalis found at the exit holes that the larvae dug out of the fruit. This behavior has been described by Roth et al. (1978) and Thompson et al. (1983) in Lixophaga diatraeae (Townsend), a very similar species, and is likely the same in L. puscolulo. After the host N. elegantalis are infested, the L. puscolulo progressively consume their internal tissues. During this process, the parasitized larvae of N. elegantalis abandon the fruit in search of shelter, such as in flowers or under dried leaves, where they finally die as a consequence of parasitism. It is known that the larvae of the last instar of N. elegantalis are parasitized before they leave the fruit, because we collected infested fruits (in the field these are recognized by the presence of excrement in the exit holes) which were taken to the laboratory and put into rearing cages with paper towels as substrate for pupation. We made periodic observations and found parasitized larvae within the paper towel material. The present study describes this new species of Lixophaga found parasitizing the larvae of the tomato fruit borer, N. elegantalis, in Colombia, with images of the parasitized 5th instar larva of N. elegantalis, and the prepupa, puparium, adult male and female, and the male terminalia of L. puscolulo.

Material and methods

Terminology follows McAlpine (1981). The description follows the format used by Byun and Han (2011), including several morphological ratios: frons-head ratio (frons width/head width); eye ratio (shortest eye diameter/ longest eye diameter); arista-antenna ratio (length of arista/length of antenna excluding arista); wing-thorax ratio (wing length/thorax length). The study was conducted at Corpoica, La Selva Research Station, located at Rionegro (Department of Antioquia, Colombia) and at Universidad del Valle (Department of Valle del Cauca, Colombia). Specimens of the new species were obtained from parasitized larvae of N. elegantalis which were naturally reared from lulo fruits, S. quitoense, collected in Colombia in the municipality of Anserma, in the State of Caldas. For the description of the new species, 21 adults were selected from a total of over 500 reared during this study. Photos were taken with a digital camera Canon EOS Rebel T3i, and images were processed with the computer program Helicon Focus® (Helicon, Soft, Ltd.). Specimens of the new species are deposited at the following institutions: USNM: National Museum of Natural History, Smithsonian Institution, Washington, DC, USA; and MUSENUV: Museo de Entomología de la Universidad del Valle, Cali, Colombia. We compared the species of Lixophaga described below with 30 Nearctic and Neotropical species present in the USNM collection, including type specimens of many of these. These are listed here with an asterisk indicating those for which primary type material was examined:

Lixophaga aberrans (Townsend)* Lixophaga angusta (Townsend) [described in Actinotachina Townsend]* Lixophaga angusta (Townsend) [described in Cataphorinia Townsend]* Lixophaga aristalis (Townsend)* Lixophaga brasiliana (Townsend)* Lixophaga charapensis (Townsend)* Lixophaga clausa (Townsend)* Lixophaga croesa (Townsend)* Lixophaga diatraeae (Townsend)* Lixophaga discalis (Coquillett)* Lixophaga fascialis (Townsend)* Lixophaga fasciata Curran Lixophaga fulvescens (Townsend)* Lixophaga fumipennis (Townsend) [described in Euprospherysa Townsend]* Lixophaga fumipennis (Townsend) [described in Paraprospherysa Townsend]* Lixophaga jennei Aldrich*

A NEW LIXOPHAGA FROM COLOMBIA Zootaxa 3737 (1) © 2013 Magnolia Press · 69 Lixophaga mediocris Aldrich* Lixophaga nigribasis Curran Lixophaga opaca Reinhard Lixophaga orbitalis Aldrich* Lixophaga parva Townsend* Lixophaga plumbea Aldrich* Lixophaga plumulosa (Townsend)* Lixophaga proletaria (Townsend)* Lixophaga punctata (Townsend)* Lixophaga retiniae (Coquillett)* Lixophaga subtilis (Wulp) Lixophaga thoracica (Curran) Lixophaga unicolor (Smith) Lixophaga variabilis (Coquillett)*

Lixophaga puscolulo Carrejo & Woodley, sp. nov. (Figs. 1–10)

Diagnosis. One character state present in Lixophaga puscolulo, the absence of apical scutellar setae, is very uncommon in Lixophaga. The following species of Lixophaga, among the 30 examined, lack apical scutellar setae and can be separated from L. puscolulo by the characters given. Lixophaga angusta (Townsend) [described in Actinotachina Townsend]. One specimen of this species was examined that lacked apical scutellar setae. This species is larger than L. puscolulo, has silvery-gray pruinosity on the head, thorax, and abdomen, and its wings are infuscated with brownish coloration. Lixophaga facialis (Townsend). This species has grayish pruinosity on the head, thorax, and abdomen, and the pruinosity on abdominal tergites 3 and 4 is present on the basal 2/3 of the tergites. Lixophaga fulvescens (Townsend). In this species abdominal tergites 1–3 are extensively yellowish laterally. Lixophaga jennei Aldrich. This Nearctic species has silvery-gray pruinosity on the head, thorax, and abdomen. Only one specimen of this species was examined that lacked apical scutellar setae. Lixophaga plumbea Aldrich. This Nearctic species has silvery-gray pruinosity on the head, thorax, and abdomen. The parafacial is much wider than in L. puscolulo and the antennal arista is thickened beyond the basal half. Lixophaga variabilis Coquillett. Some, but not all, specimens of this Nearctic species lack apical scutellar setae. This species has more silvery-gray pruinosity and on tergites 3 and 4 the pruinosity is present on the basal 2/ 3 of the tergites. Description. Body (Figs. 1, 7) generally dark colored, brownish to black, with dense yellowish pruinose areas. Male. Head (Figs. 2, 4) with a frons-to-head ratio of 0.05–0.2; eye ratio 0.01–0.1; arista-antenna ratio 0.011– 0.017; compound eye bare; inner vertical setae nearly parallel to convergent, twice as long or more than outer vertical seta; ocellar setae diverging, about as long as outer vertical seta; paravertical seta present, weak, about 0.5 times the length of the outer vertical seta; postocellar setae parallel to slightly divergent, about as long as paravertical seta; lunula glabrous, shiny; fronto-orbital plate narrow, sparsely setulose lateral to frontal setae, completely covered with yellowish pruinosity; with seven to eight frontal setae, usually six present above the antennal insertion, and two below antennal insertion, all setae strong, convergent to crossed, second pair ventral to ocelli largest, parallel, about as long as inner vertical seta; proclinate orbital seta absent; frontal vitta brownish with concolorous pruinosity, about as wide as a single fronto-orbital plate, 2.2 times as long as its median width; parafacial narrow, as long as the antenna, narrowing ventrally, completely bare, pruinosity becoming more silvery ventrally; vibrissae present, convergent, as long as the length the first flagellomere; two to four supravibrissal setae and two strong subvibrissal setae; genal dilation with surface brown, covered with silvery-white pruinosity; postgena moderately developed, completely covered with light yellow pruinosity becoming more silvery ventrally, surface with light colored setulae; postocular strip yellowish pruinose dorsally, becoming silvery-white ventrally; antenna dark in color, covered with grayish pruinosity; scape 0.5 times the length of pedicel; pedicel with strong

70 · Zootaxa 3737 (1) © 2013 Magnolia Press CARREJO ET AL. setae on the anterior margin; arista about 1.8 times the length of first flagellomere, thickened on about basal 1/4; palpus clavate, yellow, with dark-colored setae, longest laterally at apex; labellum yellowish brown, with fine light- colored setae.

FIGURES 1–4. Male of Lixophaga puscolulo Carrejo & Woodley, sp. nov. 1. Dorsal view. 2. Lateral view of head and anterior part of thorax, right side. 3. Lateral view, right side. 4. Anterior view of head.

Thorax (Fig. 1) with dorsum covered almost entirely with yellow pruinosity except for some dark areas arranged as four longitudinal vittae interrupted at suture; postpronotum with three setae arranged in form of an obtuse triangle, the angle approximately 130º–145º; notopleuron with two strong setae; scutum with three presutural and three postsutural acrostical setae; three presutural and three postsutural dorsocentral setae; three presutural and three postsutural intra-alar setae; two presutural and three postsutural supra-alar setae (two strong, anterior one weak); scutellum with basal pair of setae about 1.8 times the length of the scutellum and convergent, a pair of lateral setae 1.5 times as long as the length of a subapical seta and convergent, a pair of subapical setae 2 times the length of the scutellum and divergent, apical setae absent, and a pair of discal setae 0.8 times the length of the basal seta; pleura (Fig. 3) generally covered with grayish pruinosity that is only faintly suffused with yellow; proepisterum with one seta on ventral part; proepimeron with a strong anteroventral seta; anepisternum with two to

A NEW LIXOPHAGA FROM COLOMBIA Zootaxa 3737 (1) © 2013 Magnolia Press · 71 three anterodorsal setae and a vertical row of six to eight setae along the posterior margin; anepimeron with a moderate-sized posterodorsal seta surrounded by black setulae; katepisternum with three setae arranged in a triangle (two strong dorsal setae and one weak anteroventral seta); meron with a row of four to seven setae along the posterior margin; anterior and posterior spiracles brownish; prosternum with small, sparse setae on lateral margin; legs dark brownish to black in color, with dark colored setae and setulae, posteroventral surface of front femur with silver pruinosity; fore coxa with 4–6 anteroventral setae; fore tibia with a posteroventral seta beyond middle; mid tibia with one to two anterodorsal setae (the apicalmost largest if two), one anteroventral seta, and two posterodorsal setae; hind tibia with one large and several smaller anterodorsal setae, and 1–3 posterodorsal setae; tarsal claws simple, 0.8 times as long as 5th tarsomere; wings hyaline, completely covered with fine microtrichia; costal vein interrupted near apex of Sc; fork of R2+3 and R4 +5 with two to four small setae both dorsally and ventrally, one dorsal seta being noticeably larger than others; tegula blackish brown; calypter yellowish white with pale margin and marginal fringe of hair-like setae.

FIGURES 5–8. Female of Lixophaga puscolulo Carrejo & Woodley, sp. nov. 5. Lateral view, right side. 6. Lateral view of head and anterior part of thorax, right side. 7. Dorsal view. 8. Anterior view of head.

72 · Zootaxa 3737 (1) © 2013 Magnolia Press CARREJO ET AL. FIGURES 9–10. Male terminalia of Lixophaga puscolulo Carrejo & Woodley, sp. nov. (slide mounted). 9. Lateral view of right side. 10. Posterior view of cerci and surstyli from slide mounted specimen.

Abdomen with dark integument, tergites 2–3 and sometimes 4 yellowish laterally, tergites 3–5 with basal yellow pruinose bands occupying 1/4 to 1/3, apical portions black with sparse pruinosity, appearing moderately shiny; syntergite 1+2 with longitudinal median depression not extending to posterior margin; tergites 1+2 and 3 with a pair each of median marginal and lateral marginal setae; tergites 4 and 5 with a row of evenly spaced, long marginal setae, all tergites without discal setae. Terminalia with sternite 5 widened, U-shaped, with a strong seta on the posterior lobe on each side; cercus pointed apically (Figs. 9, 10), curved medially, with a few long posteromedial setae; surstylus narrow, apical half parallel-sided, apex curved slightly posteriorly (Figs. 9, 10), with moderately dense, anterolateral fringe of short setae; epiphallus membranous and distally rounded; pregonite elongate and slender, bent anteriorly at about middle, apex sharply rounded; distiphallus partly sclerotized, shaped as in Fig. 9; epandrium unmodified (Fig. 9), setulose. Length 5.1–5.5 mm. Female. Differs from male as follows: Head (Figs. 6, 8) with two proclinate orbital setae on each side, the anterior one distinctly larger and stouter than the posterior one; palpus slightly more robust than in male. Thorax and abdomen (Figs. 5, 7) as in male except for female terminalia. Length 4.7–6.1 mm. Material examined. Holotype male: Colombia, Caldas, Anserma, Vereda el Pensil, Finca Veracruz, 05°15΄4.5″N, 75°46΄57.4″W, 1885 m, ex fifth-instar larva of Neoleucinodes elegantalis collected on fruit of Solanum quitoense, i.2011, coll. A. E. Díaz M. (MUSENUV). Paratypes: same data as holotype: 10 adult males, 10 adult females (equally divided between MUSENUV and USNM). Etymology. The species epithet “puscolulo”, a noun in apposition, comes from the Quechua word puscolulo, which is used by the indigenous people of southern Colombia and northern Ecuador to refer to Solanum quitoense Lam. Geographical distribution. Colombia: Departments of Caldas, Cauca, Cundinamarca and Valle del Cauca, in Premontane Wet Forest (bh-PM) and Premontane Very Wet Forest (bmh-PM) according to the classification of life zones by Holdridge (1967). Host. Neoleucinodes elegantalis (Guenée) (Lepidoptera: Crambidae). Biology. A larva of N. elegantalis parasitized by L. puscolulo shows black spots on the epidermis of the body as a symptom of parasitism (Figs. 11, 12). At an average temperature of 18°C and 75% RH, survival of adult females averaged 15 days (±12.3) (n=15) and that of adult males averaged 17 days (± 11.1) (n=15). Parasitized larvae of N. elegantalis had a range of 1–4 L. puscolulo larvae per host, with an average of 1.5 (n=620). The weight of a puparium (Fig. 14) averaged 0.010 g (±0.003) (n=190) (Fig. 14) and this stage lasted an average of 24 days (±3.6) (n=34). The sex ratio in adults was 1:1 (female: male), with 78% (n=518) emergence.

A NEW LIXOPHAGA FROM COLOMBIA Zootaxa 3737 (1) © 2013 Magnolia Press · 73 Associated plants. Although N. elegantalis is known to attack many plant species of the family Solanaceae, L. puscolulo was reared only from N. elegantalis larvae collected from fruits of S. quitoense and the wild solanaceous plant S. hirtum. Remarks. As noted above the genus Lixophaga is very poorly known in the Neotropical Region with no synoptic literature, so the species are very difficult to identify with any degree of certainty. We have compared L. puscolulo with all of the available species, including type material, of the Lixophaga species housed in the USNM collection and it is clearly none of those species. In order to provide a name for this potential biocontrol agent we believe that describing the species as new is appropriate even though we have not seen every described species of Lixophaga from the Neotropical Region.

FIGURES 11–14. Host and immature stages of Lixophaga puscolulo Carrejo & Woodley, sp. nov. 11. Larva of Neoleucinodes elegantalis parasitized by Lixophaga puscolulo. Dark spots are parasitism symptoms observed on the body of the host larva. 12. Third instar larva of Lixophaga puscolulo abandoning its host larva. 13. Prepupa of Lixophaga puscolulo. 14. Puparium of Lixophaga puscolulo.

Discussion

Lixophaga puscolulo regulates populations of N. elegantalis in S. quitoense orchards in Colombia. Their populations are associated exclusively with solanaceous hosts of N. elegantalis of the Leptostemonum clade (Solanaceae with spines). This parasitoid is adapted to abiotic conditions of Premontane Wet Forest (bh-PM) and Premontane Very Wet Forest (bmh-PM). It has been observed in the field that adults of L. puscolulo mate near N. elegantalis infested fruits and the are active from 9 a.m. to 5 p.m. A natural parasitism rate of about 48% (n=850) has been observed on larvae of N. elegantalis in orchards of S. quitoense located in bmh-PM at the end of harvest, when farmers reduce insecticide applications for control of N. elegantalis (A.E. Diaz, personal observation). This means that it might be possible to increase natural populations of L. puscolulo through the implementation of simple practices such as modifying the insecticide application schedule, trying to spray after 5

74 · Zootaxa 3737 (1) © 2013 Magnolia Press CARREJO ET AL. p.m., applying active ingredients that act on N. elegantalis by ingestion and not by contact, recovering parasitoids by collecting infested fruits, and maintaining the parasitoid populations by conserving weed plants in the orchard that provide nectar and pollen to the populations of L. puscolulo. These strategies will help to preserve populations of this natural biological control agent.

Acknowledgments

Many thanks to the Regional Fund for Agricultural Technology (FONTAGRO) for funding this study; to research assistants Mr. Gilberto Higuinio and Mr. Manuel Hincapie (Corpoica, La Selva Research Station); to student Mrs. Bibiana Villada (Universidad Católica de Oriente (UCO), Rionegro, Antioquia) for the collecting of L. puscolulo in the field and its rearing in the laboratory, and for kindly providing the wonderful photos in Figs. 11–14; to Dr. William H. White (USDA-ARS Sugarcane Research Unit) for the loan of specimens of Lixophaga diatraeae used in this study for comparison with L. puscolulo.We are grateful to Dr. D. M. Wood for discussion about the of Lixophaga. Special thanks to Dr. Takumasa Kondo for translating the text originally written in Spanish, and for reviewing the manuscript. USDA is an equal opportunity provider and employer.

References

A.L.A.E. Asociación Latinoamericana de Entomología (1968) Catálogo de insectos de importancia económica en Colombia. Agricultura Tropical, Bogotá, 156 pp. Arnaud, P.H., Jr. (1978) A host–parasite catalog of North American Tachinidae (Diptera). United States Department of Agriculture, Miscellaneous Publication, 1319, 1–860. Byun, H.-W. & Han, H.-Y. (2011) Lixophaga Townsend (Diptera: Tachinidae: Blondeliini), a newly recorded parasitoid taxon in Korea. Journal of Asia-Pacific Entomology, 14 (1), 58–62. http://dx.doi.org/10.1016/j.aspen.2010.11.004 Capps, H.W. (1948) Status of the pyraustid moths of genus Leucinodes in the New World, with descriptions of new genera and species. Proceedings of United States National Museum, 98 (3223), 69–83. http://dx.doi.org/10.5479/si.00963801.98-3223.69 Crosskey, R.W. (1973) A conspectus of the Tachinidae (Diptera) of Australia, including keys to the supraspecific taxa and taxonomic and host catalogues. Bulletin of the British Museum (Natural History), Entomology Supplement 21, 1–221. Diaz, A.E. (2009) Caracterización morfométrica de poblaciones del perforador del fruto Neoleucinodes elegantalis (Guenée) (Lepidoptera: Crambidae) asociadas a especies solanáceas cultivadas y silvestres en Colombia. M Sc. thesis, Universidad Nacional de Colombia, Bogotá, Colombia. 206 pp. Diaz, A.E. (2010) The fruit borer, Neoleucinodes elegantalis (Guenée) (Lepidoptera: Crambidae), a pest of Neotropical solanaceous fruits. Abstract of a paper presented at the Potential Invasive Pests Workshop (Miami-Coconut Grove, Florida, USA, 2010-10-10/14). Diaz, A.E. & Brochero, H. (2012) Parasitoides asociados al perforador del fruto de las solanáceas Neoleucinodes elegantalis (Guenée) (Lepidoptera: Crambidae) en Colombia. Revista Colombiana de Entomología, 38 (1), 50–57. Gallego, F. & Vélez, R. (1992) Lista de insectos que afectan los principales cultivos, plantas forestales, animales domésticos y al hombre en Colombia. Universidad Nacional de Colombia y Sociedad Colombiana de Entomología, Medellín, Antioquia, Colombia, pp. 62–64. Guimarăes, J.H. (1977) Host-parasite and parasite-host catalogue of South American Tachinidae(Diptera). Arquivos de Zoologia, 28 (3), 1–131. Holdridge, L. (1967) Life zone ecology. Tropical Science Center, San José, Costa Rica. 206 pp. McAlpine, J.F. (1981) Morphology and terminology–adults. In: McAlpine, J.F, Peterson, B.V., Shewell, G.E., Teskey, H.J., Vockeroth, J.R. & Wood, D.M. (Coordinators), Manual of Nearctic Diptera. Vol. 1. Research Branch, Agriculture Canada Monograph, 27, pp. 9–63. Morales Valles, P., Cermell, M., Godoy, F. & Salas, B. (2003) Lista de insectos relacionados a las solanáceas ubicados en el museo de insectos de interés agrícola del CENIAP INIA. Entomotropica, 18, 193–209. Posada, L., Polania, I., López, A., Ruiz, N. & Rodríguez, D. (1981) Nuevo huésped. In: ICA (Ed.), Notas y Noticias Entomológicas. Bogotá, Cundinamarca, Colombia, pp. 1–11. Revelo, J., Viteri, P., Vásquez, W., León, J. & Gallegos, P. (2010) Manual del cultivo ecológico de la naranjilla. Instituto Nacional Autónomo de Investigaciones Agropecuarias, Quito, Ecuador, 119 pp. Roth, J.P., King, E.G. & Thompson, A.C. (1978) Host location behavior by the tachinid Lixophaga diatraeae. Environmental Entomology, 7, 794–798.

A NEW LIXOPHAGA FROM COLOMBIA Zootaxa 3737 (1) © 2013 Magnolia Press · 75 Sánchez, G. (1973) Las plagas del lulo y su control. Instituto Colombiano Agropecuario, Medellín, Boletín técnico, 25, 1–26. Shima, H. (2006) A host-parasite catalog of Tachinidae (Diptera) of Japan. Makunagi, Acta Dipterologica, Supplement 2, 1– 171. Thompson, A.C., Roth, J.P. & King, E.G. (1983) Larviposition kairomone of the tachinid Lixophaga diatraeae. Environmental Entomology, 12, 1312–1314. Viáfara, H.F., Garcia, F. & Díaz, A.E. (1999) Parasitismo natural de Neoleucinodes elegantalis (Guenée) (Lepidoptera: Pyralidae) en algunas zonas productoras de Solanáceas del Cauca y Valle del Cauca Colombia. Revista Colombiana de Entomología, 25, 151–159. Wood, D.M. (1985) A taxonomic conspectus of the Blondeliini of North and Central America and the West Indies (Diptera: Tachinidae). Memoirs of the Entomological Society of Canada, 132, 1–130. http://dx.doi.org/10.4039/entm117132fv Wood, D.M. & Zumbado, M.A. (2010) Tachinidae (tachinid flies, parasitic flies). Chapter 113. In: Brown, B.V., Borkent, A., Cumming, J.M., Wood, D.M., Woodley, N.E. & Zumbado, M.A. (Eds.), Manual of Central American Diptera. Vol. 2. NRC Research Press, Ottawa, pp. 1343–1417.

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