title: Review of Apanteles sensu stricto (Hymenoptera, Braconidae, Microgastrinae) from Area de Conservación Guanacaste, northwestern Costa Rica, with keys to all described species from Mesoamerica.
abstract: More than half a million specimens of wild-caught Lepidoptera caterpillars have been reared for their parasitoids, identified, and DNA barcoded over a period of 34 years (and ongoing) from Area de Conservación de Guanacaste (ACG), northwestern Costa Rica. This provides the world's best location-based dataset for studying the taxonomy and host relationships of caterpillar parasitoids. Among Hymenoptera, Microgastrinae (Braconidae) is the most diverse and commonly encountered parasitoid subfamily, with many hundreds of species delineated to date, almost all undescribed. Here, we reassess the limits of the genus Apanteles sensu stricto, describe 186 new species from 3,200+ parasitized caterpillars of hundreds of ACG Lepidoptera species, and provide keys to all 205 described Apanteles from Mesoamerica - including 19 previously described species in addition to the new species. The Mesoamerican Apanteles are assigned to 32 species-groups, all but two of which are newly defined. Taxonomic keys are presented in two formats: traditional dichotomous print versions and links to electronic interactive versions (software Lucid 3.5). Numerous illustrations, computer-generated descriptions, distributional information, wasp biology, and DNA barcodes (where available) are presented for every species. All morphological terms are detailed and linked to the Hymenoptera Anatomy Ontology website. DNA barcodes (a standard fragment of the cytochrome c oxidase I (COI) mitochondrial gene), information on wasp biology (host records, solitary/gregariousness of wasp larvae), ratios of morphological features, and wasp microecological distributions were used to help clarify boundaries between morphologically cryptic species within species-complexes. Because of the high accuracy of host identification for about 80% of the wasp species studied, it was possible to analyze host relationships at a regional level. The ACG species of Apanteles attack mainly species of Hesperiidae, Elachistidae and Crambidae (Lepidoptera). About 90% of the wasp species with known host records seem to be monophagous or oligophagous at some level, parasitizing just one host family and commonly, just one species of caterpillar. Only 15 species (9%) parasitize species in more than one family, and some of these cases are likely to be found to be species complexes. We have used several information sources and techniques (traditional taxonomy, molecular, software-based, biology, and geography) to accelerate the process of finding and describing these new species in a hyperdiverse group such as Apanteles. The following new taxonomic and nomenclatural acts are proposed. Four species previously considered to be Apanteles are transferred to other microgastrine genera: Dolichogenidea hedyleptae (Muesebeck, 1958), comb. n., Dolichogenidea politiventris (Muesebeck, 1958), comb. n., Rhygoplitis sanctivincenti (Ashmead, 1900), comb. n., and Illidops scutellaris (Muesebeck, 1921), comb. rev. One European species that is a secondary homonym to a Mesoamerican species is removed from Apanteles and transferred to another genus: Iconella albinervis (Tobias, 1964), stat. rev. The name Apanteles albinervican Shenefelt, 1972, is an invalid replacement name for Apanteles albinervis (Cameron, 1904), stat. rev., and thus the later name is reinstated as valid. The following 186 species, all in Apanteles and all authored by Fernández-Triana, are described as species nova: adelinamoralesae, adrianachavarriae, adrianaguilarae, adrianguadamuzi, aichagirardae, aidalopezae, albanjimenezi, alejandromasisi, alejandromorai, minorcarmonai, alvarougaldei, federicomatarritai, anabellecordobae, rostermoragai, anamarencoae, anamartinesae, anapiedrae, anariasae, andreacalvoae, angelsolisi, arielopezi, bernardoespinozai, bernyapui, bettymarchenae, bienvenidachavarriae, calixtomoragai, carloscastilloi, carlosguadamuzi, eliethcantillanoae, carlosrodriguezi, carlosviquezi, carloszunigai, carolinacanoae, christianzunigai, cinthiabarrantesae, ciriloumanai, cristianalemani, cynthiacorderoae, deifiliadavilae, dickyui, didiguadamuzi, diegoalpizari, diegotorresi, diniamartinezae, duniagarciae, duvalierbricenoi, edgarjimenezi, edithlopezae, eduardoramirezi, edwinapui, eldarayae, erickduartei, esthercentenoae, eugeniaphilipsae, eulogiosequeira, felipechavarriai, felixcarmonai, fernandochavarriai, flormoralesae, franciscopizarroi, franciscoramirezi, freddyquesadai, freddysalazari, gabrielagutierrezae, garygibsoni, gerardobandoi, gerardosandovali, gladysrojasae, glenriverai, gloriasihezarae, guadaluperodriguezae, guillermopereirai, juanmatai, harryramirezi, hectorsolisi, humbertolopezi, inesolisae, irenecarrilloae, isaacbermudezi, isidrochaconi, isidrovillegasi, ivonnetranae, jairomoyai, javiercontrerasi, javierobandoi, javiersihezari, jesusbrenesi, jesusugaldei, jimmychevezi, johanvargasi, jorgecortesi, jorgehernandezi, josecalvoi, josecortesi, josediazi, josejaramilloi, josemonteroi, joseperezi, joserasi, juanapui, juancarrilloi, juangazoi, juanhernandezi, juanlopezi, juanvictori, juliodiazi, juniorlopezi, keineraragoni, laurahuberae, laurenmoralesae, leninguadamuzi, leonelgarayi, lilliammenae, lisabearssae, luciariosae, luisbrizuelai, luiscanalesi, luiscantillanoi, luisgarciai, luisgaritai, luishernandezi, luislopezi, luisvargasi, manuelarayai, manuelpereirai, manuelriosi, manuelzumbadoi, marcobustosi, marcogonzalezi, marcovenicioi, mariachavarriae mariaguevarae, marialuisariasae, mariamendezae, marianopereirai, mariatorrentesae, sigifredomarini, marisolarroyoae, marisolnavarroae, marvinmendozai, mauriciogurdiani, milenagutierrezae, monicachavarriae, oscarchavesi, osvaldoespinozai, pablotranai, pabloumanai, pablovasquezi, paulaixcamparijae, luzmariaromeroae, petronariosae, randallgarciai, randallmartinezi, raulacevedoi, raulsolorsanoi, wadyobandoi, ricardocaleroi, robertmontanoi, robertoespinozai, robertovargasi, rodrigogamezi, rogerblancoi, rolandoramosi, rolandovegai, ronaldcastroi, ronaldgutierrezi, ronaldmurilloi, ronaldnavarroi, ronaldquirosi, ronaldzunigai, rosibelelizondoae, ruthfrancoae, sergiocascantei, sergioriosi, tiboshartae, vannesabrenesae, minornavarroi, victorbarrantesi, waldymedinai, wilbertharayai, williamcamposi, yeissonchavesi, yilbertalvaradoi, yolandarojasae, hazelcambroneroae, zeneidabolanosae.
authors.affiliation: Department of Integrative Biology and the Biodiversity Institute of Ontario, University of Guelph, Guelph, ON N1G 2W1 Canada ; Canadian National Collection of Insects, 960 Carling Ave., Ottawa, ON K1A 0C6 Canada.
authors.forename: Jose L
authors.initials: JL
authors.lastname: Fernández-Triana
authors.affiliation: Department of Entomology, University of Illinois, Urbana, IL 61801 USA.
authors.forename: James B
authors.initials: JB
authors.lastname: Whitfield
authors.affiliation: Dept. of Natural Sciences, Univerity of Virginia's College at Wise, Wise, VA 24293 USA.
authors.forename: Josephine J
authors.initials: JJ
authors.lastname: Rodriguez
authors.affiliation: Department of Integrative Biology and the Biodiversity Institute of Ontario, University of Guelph, Guelph, ON N1G 2W1 Canada.
authors.forename: M Alex
authors.initials: MA
authors.lastname: Smith
authors.affiliation: Department of Biology, University of Pennsylvania, Philadelphia, PA 19104-6018 USA.
authors.forename: Daniel H
authors.initials: DH
authors.lastname: Janzen
authors.affiliation: Department of Biology, University of Pennsylvania, Philadelphia, PA 19104-6018 USA.
authors.forename: Winnie D
authors.initials: WD
authors.lastname: Hallwachs
authors.affiliation: Department of Integrative Biology and the Biodiversity Institute of Ontario, University of Guelph, Guelph, ON N1G 2W1 Canada.
authors.forename: Mehrdad
authors.initials: M
authors.lastname: Hajibabaei
authors.affiliation: Department of Entomology, National Museum of Natural History, Smithsonian Institution, P.O.Box37012, MRC127, Washington, DC 20013-7012 USA.
authors.forename: John M
authors.initials: JM
authors.lastname: Burns
authors.affiliation: Systematic Entomology Laboratory, USDA, c/o National Museum of Natural History, P.O. Box 37012, Washington, DC 20013-7012, USA.
authors.forename: M Alma
authors.initials: MA
authors.lastname: Solis
authors.affiliation: Systematic Entomology Laboratory, USDA, c/o National Museum of Natural History, P.O. Box 37012, Washington, DC 20013-7012, USA.
authors.forename: John
authors.initials: J
authors.lastname: Brown
authors.affiliation: Canadian National Collection of Insects, 960 Carling Ave., Ottawa, ON K1A 0C6 Canada.
authors.forename: Sophie
authors.initials: S
authors.lastname: Cardinal
authors.affiliation: Department of Integrative Biology and the Biodiversity Institute of Ontario, University of Guelph, Guelph, ON N1G 2W1 Canada.
authors.forename: Henri
authors.initials: H
authors.lastname: Goulet
authors.affiliation: Department of Integrative Biology and the Biodiversity Institute of Ontario, University of Guelph, Guelph, ON N1G 2W1 Canada.
authors.forename: Paul D N
authors.initials: PD
authors.lastname: Hebert
keywords: Apanteles; Area de Conservación Guanacaste; Braconidae; Costa Rica; DNA barcoding; Hymenoptera Anatomy Ontology website; Lepidoptera; Lucid software; Malaise traps; Mesoamerica; Microgastrinae; caterpillar rearing; parasitoid biology; taxonomy; tropical biodiversity