Synonymy of Gerstaecker with Schiner (Diptera: ) Author(s): Allen L. Norrbom, Norman B. Barr, Peter Kerr, Ximo Mengual, Norma Nolazco, Erick J. Rodriguez, Gary J. Steck, Bruce D. Sutton, Keiko Uramoto and Roberto A. Zucchi Source: Proceedings of the Entomological Society of Washington, 120(4):834-841. Published By: Entomological Society of Washington https://doi.org/10.4289/0013-8797.120.4.834 URL: http://www.bioone.org/doi/full/10.4289/0013-8797.120.4.834

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SYNONYMY OF TOXOTRYPANA GERSTAECKER WITH ANASTREPHA SCHINER (DIPTERA: TEPHRITIDAE)

ALLEN L. NORRBOM,NORMAN B. BARR,PETER KERR,XIMO MENGUAL, NORMA NOLAZCO,ERICK J. RODRIGUEZ,GARY J. STECK,BRUCE D. SUTTON,KEIKO URAMOTO, AND ROBERTO A. ZUCCHI

(ALN) Systematic Entomology Laboratory, USDA, ARS, c/o Smithsonian Institution, P.O. Box 37012, MRC 168, Washington, DC 20013-7012, USA (e-mail: [email protected]); (NBB) Center for Plant Health Science and Technology Mission Laboratory, USDA-APHIS, Moore Air Base, 22675 N. Moorefield Rd, Edinburg, TX 78541, USA (e-mail: [email protected]. gov); (PK) California State Collection of , Plant Pest Diagnostics Branch, California Department of Food and Agriculture, 3294 Meadowview Rd., Sacramento, CA, 95832–1448, USA (e-mail: [email protected]); (XM) Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany (email: [email protected]); (NN) Centro de Diagno´stico de Sanidad Vegetal, Servicio Nacional de Sanidad Agraria, Av. La Molina 1915, La Molina, Peru´ (e-mail: [email protected]); (EJR) University of Florida; and Division of Plant Industry, Florida Department of Agriculture and Consumer Services, Gainesville, Florida 32608, USA (e-mail: erick.rodriguez@ufl.edu); (GJS) Division of Plant Industry, Florida Department of Agriculture and Consumer Services, 1911 SW 34th St., Gainesville, Florida 32608, USA (e-mail: [email protected]); (BDS) Research Associate, Department of Entomology, Smithsonian Institution, P.O. Box 37012, MRC 168, Washington, DC 20013–7012, USA (e-mail: [email protected]); (KU, RAZ) Escola Superior de Agricultura Luiz de Queiroz, Universidade de Sa˜o Paulo, Caixa Postal 9, 13418-900, Piracicaba, SP, Brazil (e-mails: [email protected]; [email protected])

Abstract.— Based on a recent phylogenetic analysis, Toxotrypana Gerstaecker (1860) is here placed in synonymy with Anastrepha Schiner (1868). Although Toxotrypana is the senior name, Anastrepha is recognized as valid because of the much greater economic significance of its species. Changing the names of the many pest species of Anastrepha would cause major nomenclatural instability and con- fusion in the applied literature, therefore a proposal to the International Commission on Zoological Nomenclature has been submitted to validate this usage. The following nomenclatural changes are proposed: Anastrepha australis (Blanchard 1960), new combination, Anastrepha curvicauda (Gerstaecker 1860), new combination, Anastrepha littoralis (Blanchard 1960), new combination, Anastrepha nigra (Blanchard 1960), new combination, Anastrepha picciola (Blanchard 1960), new combination, Anastrepha proseni (Blanchard 1960), new combination, Anas- trepha recurcauda (Tigrero 1992), new combination (all transferred from Toxo- trypana), and Anastrepha nigrina Norrbom, new name for Anastrepha nigra VOLUME 120, NUMBER 4 835

Norrbom and Korytkowski (2009). Some changes to the species group classification within Anastrepha are also discussed. Key Words: fruit , , nomenclature, new name, new combination DOI: 10.4289/0013-8797.120.4.834

Toxotrypana Gerstaecker (1860) is clade of Anastrepha, and this lineage is a small genus of fruit flies (Diptera: sister to the tripunctata group of Anas- Tephritidae) restricted to the New World trepha (taxa sensu Norrbom et al. tropics and subtropics. Its species are 2012). presumed to be mimics of vespid wasps The purpose of this publication is to due to their body shape, bright yellow formally recognize the synonymy of and brown coloration, reduced setation, Anastrepha and Toxotrypana and to and their wing patterns with a broad, make changes to the names of the spe- complete costal band. Despite their dis- cies impacted by this synonymy. Some tinctive external appearance, the species changes to the species group classifica- of Toxotrypana have been known for tion within Anastrepha based on the some time to be closely related to results of the Mengual et al. (2017) Anastrepha Schiner (1868) on the basis analysis are also discussed. of genitalic characters (Norrbom et al. 1999b). Anastrepha and Toxotrypana together MATERIALS AND METHODS form a well-defined monophyletic group We follow the morphological termi- supported by both morphological nology of White et al. (1999), except for (Norrbom et al. 1999b) and molecular the wing venation, which follows that of studies (Han and McPheron 1997, Cumming and Wood (2017). McPheron et al. 1999, Han and Ro 2009). Toxotrypana also is clearly RESULTS AND DISCUSSION monophyletic (Norrbom et al. 1999b), Anastrepha Schiner Synonymy but the status of Anastrepha has been uncertain. Some studies suggested that it Anastrepha Schiner 1868: 263, type spe- might be paraphyletic, but were not cies Dacus serpentinus Wiedemann conclusive due to limited taxon sampling 1830, by original designation. or insufficient character evidence (McPheron Toxotrypana Gerstaecker 1860: 191, type et al. 1999, Norrbom et al. 1999b, Barr et al. species T. curvicauda Gerstaecker 2005). Recently, however, Mengual et al. 1860, by monotypy; New syno- (2017) investigated relationships within nym. the Anastrepha/Toxotrypana clade, uti- Acrotoxa Loew 1873: 227, type species lizing sequence data from six DNA re- Dacus fraterculus Wiedemann 1830, gions in a total of 150 species. Their by subsequent designation of Bezzi results conclusively demonstrate that 1909: 280. Toxotrypana arises within Anastrepha, Mikimyia Bigot 1884: xxix, type species confirming that Anastrepha as currently M. furcifera Bigot 1884 (= curvi- defined is paraphyletic. Toxotrypana is cauda Gerstaecker), by monotypy. the sister group of the cryptostrepha New synonym. 836 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Pseudodacus Hendel 1914a: 66, type commercial and subsistence crops such as species Anastrepha daciformis Bezzi citrus, guava, mango, melon, and many 1909, by original designation (as others (White and Elson-Harris 1992, subgenus of Anastrepha). Norrbom 2004). Numerous other species Pseudodacus Hendel 1914b: 13, type are minor or potential pests. Several species Anastrepha daciformis Bezzi species are invasive, including A. fra- 1909, by original designation (as terculus (Brazil-1 type in Argentina, subgenus of Anastrepha). Pre- Bolivia and Peru), A. grandis (Panama), occupied by Hendel 1914: 66. A. ludens (Panama), and A. suspensa Phobema Aldrich 1925: 7, type species (Florida) (Weems 1965, 1966; Sutton P. atrox Aldrich 1925, by original et al. 2015). designation. Despite differences in their external Lucumaphila Stone 1939: 340, type appearance, Toxotrypana and Anas- species L. sagittata Stone 1939, by trepha have been considered to be original designation. closely related on the basis of genitalic characters (Norrbom et al. 1999b). Al- Anastrepha is the largest genus of though it has been clear that together Tephritidae in the New World, including they form a monophyletic group, both nearly 300 valid species (Norrbom et al. have continued to be recognized as 1999a, 2012, 2015), with additional new genera because of the distinctiveness of species in the process of description Toxotrypana (based on numerous auta- (Norrbom et al., in prep.). Toxotrypana pomorphies) and the uncertainty of the currently includes seven valid species exact relationship of the two taxa, par- and a similar number of undescribed ticularly whether or not Anastrepha, species (Norrbom et al. 1999a, 1999b). sensu lato, is monophyletic. Both genera include agricultural pests, Based on the results of the phyloge- but Anastrepha is by far more econom- netic analysis by Mengual et al. (2017) ically important. Toxotrypana curvi- that Anastrepha, sensu lato, is para- cauda Gerstaecker, commonly known as phyletic, we here formally recognize the the fruit fly, is a major pest of synonymy of Anastrepha and Toxo- papaya (Carica papaya L.) in the circum- trypana. Although Toxotrypana has Caribbean area (White and Elson-Harris priority, Anastrepha has far greater eco- 1992, Norrbom 2004). It is invasive in nomic importance and treating the latter Florida (Weems 1969). Other species of as invalid would cause tremendous Toxotrypana are not considered to be nomenclatural instability and disrupt economically significant. scientific communication regarding Multiple species of Anastrepha, in- multiple major pest species. Thus, we cluding at least three members of the A. recognize Toxotrypana as a subjective fraterculus (Wiedemann) complex (South synonym of Anastrepha. A proposal to American fruit fly), A. grandis (Mac- the International Commission on Zoolog- quart) (South American cucurbit fruit ical Nomenclature has been submitted to fly), A. ludens (Loew) (Mexican fruit fly), validate this usage (Norrbom et al. 2018). A. obliqua (Macquart) (West Indian fruit The species previously classified in Toxo- fly), A. serpentina (Wiedemann) (sapote trypana are henceforth recognized as the fruit fly), A. striata Schiner (guava fruit curvicauda species group of Anastrepha. fly), and A. suspensa (Loew) (Caribbean The nigra subgroup of the robusta fruit fly) are major pests, impacting group was recognized by Norrbom and VOLUME 120, NUMBER 4 837

Korytkowski (2009) to include A. nigrina group. Norrbom et al. (2014) added A. Norrbom (as A. nigra Norrbom and tehuacana Norrbom to the latter group, Korytkowski; see Nomenclatural Changes) and Norrbom et al. (2015) added A. and A. partita Norrbom and Korytkowski. disjuncta Norrbom to the former This clade was erroneously omitted from group. the Norrbom et al. (2012) species group The cryptostrepha group, as originally classification. Based on the results of the proposed, was based on similarities in Mengual et al. (2017) analysis, it is rec- the shape of the male surstyli (character ognized here as a separate species group polarity uncertain) and aculeus tip (like most of the subgroups of the robusta (apomorphic, but not unique), reduced group), and it is here renamed the nigrina scutal microtrichia (apomorphic, but not group to reflect the change in name of its unique), and weak apical curvature of namesake included species. vein M1 (plesiomorphic). Results of the The cryptostrepha group was pro- Mengual et al. (2017) analysis indicate posed by Norrbom and Kim (1988) and that the cryptostrepha group, sensu lato, Norrbom et al. (1999b) to include is not monophyletic and that these mor- Anastrepha cordata Aldrich, A. cryptos- phological characters are homoplasious. trepha Hendel, A. margarita Caraballo, A. Their analysis supports the Norrbom panamensis Greene, A. tripunctata Wulp, et al. (2012) classification of these spe- and A. zeteki Greene. It was considered cies. The lone representative of the tri- asubgroupoftherobusta group by punctata group, A. tehuacana, is placed Norrbom and Korytkowski (2009) and in the Mengual et al. (2017) analysis as Norrbom et al. (2012), and restricted to A. the sister group of the cryptostrepha cordata and A. cryptostrepha,plustwo group, sensu stricto, plus the curvicauda new species, A. cryptostrephoides Norrbom group (former Toxotrypana), but the and Korytkowski and A. disticrux Norrbom panamensis group is clustered with A. and Korytkowski. The three species punctata Hendel, rather distant from this treated by Mengual et al. (2017) formed clade. a monophyletic group apart from the Nomenclatural Changes other subgroups of the robusta group and the subgroup was therefore recognized The following nomenclatural changes as a separate species group. are necessary because of the synonymy The other species originally included of Toxotrypana with Anastrepha, in- in the cryptostrepha group were placed cluding transfer of the seven valid spe- in new species groups by Norrbom et al. cies originally described in Toxotrypana (2012). Although no explanation was which are hereby transferred to Anas- provided with that list, the division of the trepha, and one replacement name: species followed the hypotheses of re- lationship within the original cryptos- Anastrepha australis (Blanchard), trepha group discussed by Norrbom new combination et al. (1999b). Anastrepha margarita, A. panamensis, and A. zeteki were placed in Toxotrypana australis Blanchard 1960: the panamensis group, whereas A. tri- 34. Syntype females (Museo Ar- punctata was included with A. maya gentino de Ciencias Naturales). Herna´ndez-Ortiz and A. relicta Herna´ndez- Type locality - Argentina: Tucuma´n: Ortiz (the “two n. spp. nr. tripunctata”of Tucuma´n. Distribution - Argentina Norrbom et al. 1999b) in the tripunctata (Buenos Aires, Corrientes, Santa Fe, 838 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Santiago del Estero, Tucuma´n), Bolivia de Ciencias Naturales). Type local- (Tarija), Brazil (Rio Grande do Sul). ity - Argentina: Entre Rios: Con- cordia. Distribution - Argentina (Co´r- doba, Entre Rios, Jujuy, Santa Fe), Anastrepha curvicauda (Gerstaecker), Bolivia (Cochabamba). new combination Toxotrypana pseudopicciola Blanchard Gerstaecker 1960: 43. Holotype female (Museo 1860: 194. Type female (Museum Argentino de Ciencias Naturales). fur Naturkunde der Humboldt Uni- Type locality - Argentina: Co´rdoba: versitat zu Berlin). Type locality - Co´rdoba. Synonymy (Norrbom et al. Virgin Islands: Insula St. Jean [St. 1999b: 338). John]. Distribution - Bahamas, Belize, Colombia, Costa Rica, Cuba, Dominican Republic, Neth- Anastrepha nigrina Norrbom, erlands Antilles, El Salvador, Gua- new name temala, Honduras, Me´xico (Chiapas, Anastrepha nigra Norrbom and Korytkowski Morelos, Oaxaca, Tamaulipas, Vera- 2009: 37. Holotype female (Na- cruz), Panama´, Puerto Rico, Trinidad, tional Museum of Natural History, USA (southern Texas), Venezuela, Smithsonian Institution). Type Virgin Islands; introduced USA locality - Panama´: Panama´: Parque (Florida). Nacional Chagres, Altos de Pacora Mikimyia furcifera Bigot 1884: xxix. [9°15928"N 79°21924"W], Lote H4. Holotype male (University Mu- Preoccupied by nigra Blanchard 1960: seum, Oxford), Type locality: Brazil 42. Distribution - Panama´.Etymology- [probably erroneous]. The name is a Latin adjective in ref- Toxotrypana fairbatesi Munro 1984: erence to the dark wing pattern. 160. Holotype male (South African National Collection of ). Type locality - Cuba: Soledad, Sta. Clara. Anastrepha picciola (Blanchard), Synonymy (Steyskal 1986: 114). new combination Toxotrypana picciola Blanchard 1960: 40. Holotype female (Museo Argen- Anastrepha littoralis (Blanchard), tino de Ciencias Naturales). Type lo- new combination cality - Argentina: Tucuma´n: Estacio´n Toxotrypana littoralis Blanchard 1960: Experimental Agrı´cola de Tucuma´n. 36. Holotype female (Museo Ar- Distribution - Argentina (Tucuma´n). gentino de Ciencias Naturales). Type locality - Argentina: Corrientes: Cor- rientes. Distribution - Argentina, Gua- Anastrepha proseni (Blanchard), temala, Peru (Cusco), Venezuela. new combination Toxotrypana proseni Blanchard 1960: 38. Holotype female (Museo Ar- Anastrepha nigra (Blanchard), gentino de Ciencias Naturales). new combination Type locality - Argentina: Buenos Toxotrypana nigra Blanchard 1960: 42. Aires: Punta Lara. Distribution - Holotype female (Museo Argentino Argentina (Buenos Aires, Jujuy). VOLUME 120, NUMBER 4 839

Anastrepha recurcauda (Tigrero), section, cell cua with large posteroapical new combination lobe. Male terminalia: Surstyli relatively Toxotrypana recurcauda Tigrero 1992: short, lateral surstylus without anterior 102. Holotype female (Pontificia or posterior lobes (autapomorphy?). Universidad Cato´lica del Ecuador). Subepandrial sclerite with strongly Type locality - Ecuador: Loja: Mal- sclerotized secondary connection. Glans acatos, Nangora, 79°6’W 04°13’S, slender and weakly sclerotized medially, 1,600 m. Additional description with T-shaped apical sclerite (autapo- and type data (Tigrero 1998: 6). morphy), or absent (synapomorphy of Distribution - Ecuador. daciformis and dentata groups). Generic Diagnosis for Anastrepha Female terminalia: Oviscape with lateral, flange-like lobes basally (auta- The best diagnostic characters for pomorphy), tubular, often elongate, oc- Anastrepha as here recognized are gen- casionally strongly dorsally arched. italic, but most species can be recog- Eversible membrane enlarged basally nized by their wing pattern and the other and bearing group of enlarged dorsal external characters listed below. denticles (autapomorphy; denticles sec- Body usually predominantly orange, ondarily reduced in dentata group, but occasionally predominantly brown or still larger than in this area in most other with brown markings, usually with white tephritids, and the basal area still en- or yellow markings, although sometimes larged). Aculeus tip with 3 pairs of short not well differentiated (typically in- sensilla ventrally, none extended beyond cluding at least postpronotal lobe, post- lateral margin; lateral margin serrate or sutural sublateral vitta on scutum, entire. scutellum at least apically, dorsal margin of anepisternum, posterior margins of abdominal tergites, and often medial ACKNOWLEDGMENTS vitta on scutum). We sincerely thank Neal Evenhuis, Thorax: Dorsocentral seta aligned Thomas Pape, and two anonymous re- closer to postalar seta than to postsutural viewers for their comments on previous supra-alar seta or posterior to postalar drafts of the manuscript. Mention of seta, rarely absent (some species of trade names or commercial products in curvicauda group). this publication is solely for the purpose Wing: Pattern usually consisting of C-, of providing specific information and S- and V-bands variously connected or does not imply recommendation or en- separated, occasionally instead with dorsement. USDA is an equal opportu- only uninterrupted costal band and cu- nity provider and employer. bital streak (wasp mimic pattern), rarely A. api- with other type of pattern (e.g., Literature Cited cata Norrbom and Korytkowski, A. ob- scura Aldrich, male A. lutea Stone). Aldrich, J. M. 1925. New Diptera or two-winged flies in the United States National Museum. Vein M1 anteriorly curved on distal half of last segment (distal to crossvein dm-m), Proceedings of the United States National Museum 66 (18) [= No. 2555]: 1‒36 pp. usually strongly curved apically and Barr, N. B., L. Cui, and B. A. McPheron. 2005. meeting costa without visible angle. Molecular systematics and sequence analysis Vein CuA with strong bend in middle of the nuclear gene period in the genus 840 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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