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ZooKeys 383: 1–565, doi: 10.3897/zookeys.383.6418
Review of Apanteles sensu stricto (Hymenoptera, Braconidae, Microgastrinae) from Area de Conservación Guanacaste, northwestern Costa Rica, with keys to all described species from Mesoamerica
Jose L. Fernández-Triana 1,2,†, James B. Whitfield 3,‡, Josephine J. Rodriguez 4,§, M. Alex Smith 1,|, Daniel H. Janzen 5,¶, Winnie D. Hallwachs 5,#, Mehrdad Hajibabaei 1,††, John M. Burns 6,‡‡, M. Alma Solis 7,§§, John Brown 7,||, Sophie Cardinal 2,¶¶, Henri Goulet 1,##, Paul D. N. Hebert 1,†††
1 Department of Integrative Biology and the Biodiversity Institute of Ontario, University of Guelph, Guelph, ON N1G 2W1 Canada
2 Canadian National Collection of Insects, 960 Carling Ave., Ottawa, ON K1A 0C6 Canada
3 Department of Entomology, University of Illinois, Urbana, IL 61801 USA
4 Dept. of Natural Sciences, Univerity of Virginia’s College at Wise, Wise, VA 24293 USA
5 Department of Biology, University of Pennsylvania, Philadelphia, PA 19104-6018 USA
6 Department of Entomology, National Museum of Natural History, Smithsonian Institution, P.O.Box37012, MRC127, Washington, DC 20013-7012 USA
7 Systematic Entomology Laboratory, USDA, c/o National Museum of Natural History, P.O. Box 37012, Washington, DC 20013-7012, USA

Corresponding author: Jose L. Fernández-Triana (jftriana@uoguelph.ca)

Academic editor: K. van Achterberg

received 11 October 2013 | accepted 15 January 2014 | Published 24 February 2014
(C) 2014 Jose L. Fernández-Triana. 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.
For reference, use of the paginated PDF or printed version of this article is recommended.

Citation: Fernández-Triana JL, Whitfield JB, Rodriguez JJ, Smith MA, Janzen DH, Hallwachs W, Hajibabaei M, Burns JM, Solis MA, Brown J, Cardinal S, Goulet H, Hebert PDN (2014) 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. ZooKeys 383: 1–565. doi: 10.3897/zookeys.383.6418

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.

Keywords

Apanteles, Microgastrinae, Braconidae, taxonomy, parasitoid biology, DNA barcoding, Lepidoptera, caterpillar rearing, Malaise traps, tropical biodiversity, Area de Conservación Guanacaste, Costa Rica, Mesoamerica, Lucid software, Hymenoptera Anatomy Ontology website

Introduction

The subfamily Microgastrinae (Hymenoptera: Braconidae) is one of the most speciose subfamilies of parasitoid wasps, with more than 2, 200 described species (Yu et al. 2012) and many thousands more awaiting description (Mason 1981; Rodriguez et al. 2012). Microgastrine wasps are significant in biological control because they attack the larvae of most families of Lepidoptera (Whitfield 1995, 1997).

The genus Apanteles was erected by Förster (1862) to include all species of microgastrines lacking a second submarginal cell in the fore wing (from the Greek: A – without, panteles – complete, entire; referring to the “incomplete” venation, i.e., missing cell, when compared with the other genera of Microgastrinae known at the time). As the study of Apanteles progressed, it became evident that it included a huge number of species, and many attempts to subdivide the genus have been made since 1880; there are summarized in Mason (1981) and Whitfield et al. (2002). During the last 150 years more than two dozen new genera have been created as a result of those splitting efforts, but still more than one thousand described species belong to Apanteles (Yu et al. 2012), and thousands more await discovery. It is worth mention that many of these species still belong to Apanteles sensu lato, and have not yet been assigned to currently recognised genera (sensu Mason 1981).

Area de Conservación Guanacaste (ACG) is a single decentralized unit of Costa Rica’s Ministerio del Ambiente, Energia (MINAE; Ministry of Environment and Energy) covering about 2% of Costa Rica in its northwestern corner, slightly south of the southeastern border of Nicaragua (http://www.acguanacaste.ac.cr). Comprising ~1, 200 km2 of terrestrial habitat (centered at 10.8 latitude, -85.6 longitude), it is a swath from Pacific coastal mangroves across lowland dry forest (dry season deciduous), up the slopes of three volcanoes to cloud forest (1400–2000 m), and down into Caribbean lowland (90 m) rain forest. It is only 85 km from east to west, yet contains portions of eight Holdridge Life Zones within mosaics of them, some as small as 5 km in linear dimensions and 20 km2. Nearly all of the ACG lowlands have been subjected to four centuries of light to intense cultivation, logging, burning, hunting, ranching, and other forms of habitat destruction, followed by explicit protection and restoration beginning in 1971 and intensifying after 1985 (Janzen 1988, 2000, 2002). The outcome is a mosaic of all imaginable ages and kinds of secondary succession intermingled with tiny to medium-sized fragments of approximations of intact forest (more intact in upper elevations than lower), as well as severe blurring and elimination of interdigitated boundaries between habitats and ecosystems (Janzen 1986-1988). All of the ACG region has also now experienced at least two decades of notable drying and increasing weather unpredictability, rendering it yet more difficult to know if the marked annual and decadal population changes are being generated by climate changes, successional changes, insularization of the ACG ecological island in the agroscape, species-by-species biological serendipity, and/or interactions among all of these (Janzen et al. 2011).

ACG has been the focus of 34+ years of inventory of wild-caught caterpillars, their food plants and their parasitoids, as described in detail in Janzen et al. (2009) and Janzen and Hallwachs (2011), and available in a rearing-by-rearing specimen-based public database at Janzen and Hallwachs (2013). The ACG is currently staffed and supported by about 180 Costa Ricans, nearly all of whom are honoured with patronynms in this paper.

Haphazardly placed Townes Malaise traps in all three major ACG terrestrial ecosystems have yielded another set of ACG Apanteles species, many of which have not yet been reared and are included here (and are so indicated as distinct from the species that have been reared, many of which have not yet been encountered by Malaise-trapping).

The rearing results have been complemented since 2003 by extensive DNA barcoding of one or more voucher specimens from each rearing, past and present (Janzen and Hallwachs 2011). This has provided an additional layer of data to study the ACG species of caterpillars, parasitoids, and food plants (e.g., Smith et al. 2006, 2007, 2008; Whitfield et al. 2012; Janzen et al. 2011, 2012).

DNA barcoding uses a short standardized region of the mitochondrial gene cytochrome c oxidase (COI) as a key character for species-level identification and discovery (Floyd et al. 2002, Hebert et al. 2003a and b, Janzen et al. 2009, Smith et al. 2006, 2007, 2008). Interspecific barcode variation can be used as part of a suite of characters for the discovery and description of new species (e.g., Hebert et al. 2004, Burns et al. 2008, Fisher and Smith 2008, Fernández-Triana 2010), and can speed the rate of taxonomic research by flagging otherwise cryptic diversity (e.g., Janzen et al. 2009, Fisher and Smith 2008, Smith and Fisher 2009, Smith et al. 2008). DNA barcoding has been extensively used in biodiversity and taxonomic studies of Microgastrinae during the past five years (e.g., Smith et al. 2008 and 2013, Janzen et al. 2009, Fernández-Triana 2010, Fernández-Triana et al. 2011, Rodriguez et al. 2012, Whitfield et al. 2012, Fernández-Triana et al. 2013).

Taxonomic studies of ACG Microgastrinae have been published elsewhere (e.g., Valerio et al. 2005, Grinter et al. 2009, Smith et al. 2008, Valerio et al. 2009, Janzen and Hallwachs 2011, Janzen et al. 2009, Whitfield et al. 2012, Arias-Penna et al. 2013, Fernández-Triana et al. 2013). However, the ACG species of Apanteles sensu stricto have never been treated in a taxonomic review.

The combination of this comprehensive inventory with the richness of biological, ecological and DNA barcoding data, allowed us to engage in the taxonomic study of ACG Apanteles as a whole, and within the context of the other hundreds of species of ACG Microgastrinae. In doing so, we also revised all 19 of the previously described Apanteles sensu stricto known from Mesoamerica and incorporate them here. However, no effort was made to study specimens representing undescribed species from areas outside ACG, areas that will certainly contain hundreds of other species of Apanteles as well as many of those in ACG. We hope that this study will be a foundation upon which future studies of tropical Apanteles and other microgastrine genera can be based.

Methods

In this study, Mesoamerica is defined as the region from (and including) Mexico through Panama, and all the Caribbean islands, following Gauld (1988).

We studied 4, 100+ specimens from 3, 200+ individual caterpillar rearings, and 2, 000+ DNA sequences (usually one sequence per rearing event) of Apanteles from ACG. Ecological, biological and distribution data for all of these records can be accessed at http://janzen.sas.upenn.edu/caterpillars/database.lasso (Janzen and Hallwachs 2013) by searching on the “DHJPARxxxxxxx” voucher code of the wasp, or the “yy-SRNP-xxxxx” voucher codes of the caterpillar. If a DHJPARxxxxxxx voucher code is cited, it is for a single specimen. If a yy-SRNP-xxxxx voucher code is cited, it is for 1 to N specimens reared from a single caterpillar and which are presumed siblings, but have not been individually vouchered, whether point-mounted or remaining preserved in ethanol. All holotypes bear a DHJPARxxxxxxx unique voucher code (and if there was more than one specimen in that rearing from that one caterpillar, all of them will bear the same yy-SRNP-xxxxx code). In this paper we refer to these voucher codes as “ACG database codes” when providing specimen details in the taxonomic treatment of species. In the case that a set of specimens reared from one individual caterpillar was not DNA barcoded, the vial containing those specimens has only the yy-SRNP-xxxxx code, while an individual wasp that has been barcoded from that sample bears both the SRNP code and the DHJPAR code. Each barcoded specimen also has an accession code from the Barcode of Life Data System (BOLD) and GenBank.

Type material for most of the 19 previously described Mesoamerican species was borrowed for study. However, no molecular data is available for any of those holotypes. It will not be surprising if some of their names are found to encompass complexes of species. Some members of such complexes may be some of the ACG species described here, but it would be premature to even speculate about that.

The following acronyms are used:

BMNH The Natural History Museum, London, United Kingdom

CNC Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa, Canada

INHS Illinois Natural History Survey, Champaign, Illinois, United States

INBio Instituto Nacional de Biodiversidad, Santo Domingo de Heredia, Costa Rica

NMNH National Museum of Natural History, the Smithsonian Institution, Washington DC, United States

Morphological terms and measurements of structures are mostly as used by Mason (1981), Huber and Sharkey (1993), Sharkey and Wharton (1997), Whitfield (1997), and Valerio et al. (2009). However, we also incorporated a recent, comprehensive morphological treatment of Opiinae (Braconidae) by Karlsson and Ronquist (2012), which is part of a wider effort to standardize and homologize morphological terms and definitions across the order Hymenoptera, the Hymenoptera Anatomy Ontology (HAO) project (Yoder et al. 2010, Seltmann et al. 2012). As a result of adopting most HAO preferred terms (but see exceptions below), some of the morphological terms we apply have never been used in taxonomic papers treating Microgastrinae.

Karlsson and Ronquist (2012) named and numbered the first metasomal segment as “abdominal tergum/sternum 2”. Usually, that segment has been called (and numbered) “metasomal tergum/sternum 1” (Mason 1981, Whitfield 1997, 2006). Though both approaches are correct, we use “metasomal tergum/sternum 1” because we consider it is clearer and facilitates the counting of metasomal segments (as in Fig. 207). The same applies to its associate sclerites (mediotergites and laterotergites).

We considered that the “preferred label” (i.e., name) provided in the HAO website for “mesoscutellar arm” was better than the corresponding term, “posterior bar of mesoscutellum”, used by Karlsson and Ronquist (2012).

The terms “mesoscutellar trough” and “mesoscutellar arm” (Fig. 206) have been used extensively in taxonomy of Microgastrinae (e.g., Mason 1981, and many subsequent papers), usually under the name of “lateral face of scutellum”. Karlsson and Ronquist (2012) did not provide much detail for those areas because the two species of Opiinae they studied are relatively simple and non-differentiated in that body region.

We also calculated and compared many ratios between linear dimensions of structures (morphometric taxonomy), a common practice in the taxonomy of many groups of parasitoid wasps (Baur and Leuenberger 2011, and references cited there). However, most of the ratios presented here have not been used previously in Microgastrinae taxonomy.

To facilitate understanding of the traits and ratios, a detailed account of every morphological structure and measurement used in this study is provided in Appendix 1, including links to the HAO website and references to terms that have been commonly used previously in Microgastrinae taxonomy. The most important morphological characters used in this study are illustrated in Figs 206209.

Throughout the text, especially in the keys, “body length” refers to the length of the anatomical line that is median and extends between the anteriormost point of the head and the posteriormost point of the metasoma (excluding ovipositor and ovipositor sheaths). “Fore wing length” refers to the length of the anatomical line that extends between the median margin of the first axillary sclerite and the distalmost point of the wing blade (Appendix 1).

The measurement of variables must be done as uniformly as possible, and special care must be taken when choosing the end points of any structure. It is also advisable to measure at the highest possible magnification to minimize errors. Some measurements that are particularly error-prone are discussed further in Appendix 1.

Throughout the keys the following acronyms are used for morphological terms: T1, T2, T3 (mediotergite 1, 2, 3). Whenever there is a “(N = a number)”, e.g., “(N = 4)” after a species name, it refers to the number of specimens studied morphologically for that species. It is only provided when the available number of specimens was less than 5.

Molecular analysis has revealed a large number of morphologically cryptic species, often possessing very subtle morphological differences that we found to correlate with ecological and host data. Certain features differ just slightly between species, and there may be overlap of values between individual specimens of different but very similar species. We studied as many specimens as were available. Our definition of a “species” is a postulated biological unit that differs from other species in its morphology (however subtle), COI barcode, and host use, and presumably represents a distinct breeding population. In the few cases where what we consider to be a species differs only in barcode and/or host, we indicate this. All the species in ACG are essentially fully sympatric to parapatric (the case when two ecosystems intergrade).

The dichotomous keys were built to accommodate, as much as possible, what appear to be potential natural groups, based on morphology, biology (host data), and DNA barcoding. However, in such a large assemblage of species there is likely to be considerable homoplasy and thus in some couplets we had to use logical characters (e.g., “if”, “then”, “and”, “or”, “and/or”). Those words are shown in bold and italic throughout the keys, to be explicit that in those cases more than one character system has to be considered.

The species descriptions are based on the holotype female, and we consider their DNA barcodes to be definitive when available (actual barcodes are available through the BOLD web site at http://www.boldsystems.org); when other specimens are available, their data are included to provide some idea of intraspecific variation. When the holotype was not examined or was lacking some body parts (some old holotypes), other specimens were used to complete the descriptions, and details were explained in the “Comments” section.

Males of Microgastrinae are difficult to key out under the present knowledge of the subfamily (Whitfield 1997, 2006), and may be difficult or impossible to identify unless associated with sibling females from the same host caterpillar, DNA barcodes, or host data. Thus, the keys are only intended for female specimens and use many characters only found in that sex (e.g., length of ovipositor sheaths).

Non-morphological characters are also provided whenever available, e.g., host species, whether there are one or many larvae per host caterpillar, microhabitat, microgeographical distribution, and molecular differences in the DNA barcode region, that may serve as diagnostic characters. Sometimes those features are included in brackets at the end of the corresponding couplet, intended as supplementary information that can help the user to correctly identify specimens. They are best not separated from the morphological features provided. However, in future, practical and routine identification may often depend heavily on these other traits because they are easier to assess than the morphology of wasps 1–5 mm in length.

Lucid 3.5.4 (http://www.lucidcentral.com/) software was used to automatically generate descriptions of the species and to prepare Lucid identification keys. A dataset of 41 characters and 239 character-states was used to provide uniform description formats for all species treated (except for the leucostigmus species-group, see next paragraph). The description format includes one sentence per character, with the character mentioned first and the character-state following after a colon, e.g., “Tarsal claws: simple”. Whenever a species scored more than one character-state, the description included all of the pertaining character-states separated by “or”, e.g., “Tarsal claws: simple or simple with single basal spine-like seta”. Whenever a character-state was coded as uncertain due to poor condition of the specimen(s), the description includes the details of the character-state as best assessed, followed by a question mark, e.g., “Tarsal claws: simple (?)”. Sometimes a character could not be coded due to missing body parts in the available specimens; in such instances the feature was left out of the description for that particular species.

The leucostigmus species-group was found to be exclusively composed of morphologically cryptic species (e.g., Smith et al. 2008). In this group we only provide “complete” classical descriptions (i.e., similar to other groups) for the two species previously described, Apanteles albinervis (Cameron, 1904) and Apanteles leucostigmus (Ashmead, 1900). Another 37 new species found to belong to this group are only described using a simplified set of 10 characters and 46 characters-states, and are most definitively defined by the combination of their barcodes and their hosts.

Accurate color description is a major difficulty in taxonomic works (Aguiar 2005), and often its variation within and between species can lead to confusion. In Microgastrinae, as in many other groups, the color pattern of body regions seems to be more important and taxonomically informative than the definition of the color per se. Accordingly, in the Lucid dataset, we used a simplified convention to code color, considering it as either pale (white, light yellow, orange-yellow, light brown-yellow) or dark (dark brown, black). For details on the exact color patterns on the body, we provide extensive illustrations for every species. When describing leg color (especially metacoxa, metafemur, and metatibia), we are referring to the outer side of the leg, e.g., sensu figure 1 of Goulet and Huber (1993).

Most of the photos were taken with a Canon EOS 60D with MPE-65 lenses (aperture: 4.0, ISO: 100, CR2 format images), and a 600EX-RT Speedlight (manual) flash. The camera was mounted on a Kaiser copy stand with a Z-stepper (Stackshot) to allow for taking of multiple images. A dozen species were photographed with a Keyence VHX-1000 Digital Microscope, using a lens with a range of 13–130 ×.

Multiple images through the focal plane were taken of a structure and these were combined to produce a single in-focus image. For the pictures taken with the Canon camera, the Zerene Stacker program (http://zerenesystems.com/cms/stacker) was used; the software associated with the Keyence System produced the focused images taken with that camera. Plates for the illustrations were prepared using Adobe Photoshop CS4.

Although the keys provided in this paper are based on morphological characters, in a few cases (mainly with species belonging to the leucostigmus group) we used molecular characters to differentiate species that are morphologically exceptionally similar to each other. In those cases we used characteristic loci in the DNA barcoding region in identification couplets. The bases are numbered from the start of the COI gene according to the reference sequence U37541 (Drosophila melanogaster). The bases noted are only diagnostic within the couplet and beneath that described split. A hypothetical example of the format used is: “A total of 11 diagnostic characters in the barcoding region: 12 A, 18 C, 22 A, 23 T, 44 C, 56 G, 120 G, 340 A, 488 T, 502 T, 601 A”. The letters A, C, G, and T correspond to adenine, cytosine, guanine, and thymine respectively.

DNA barcodes for all ACG inventory Microgastrinae (Appendix 2) were obtained using DNA extracts prepared from single legs using a glass fibre protocol (Ivanova et al. 2006). Extracts were re-suspended in 30 μl of dH2O, and a 658-bp region near the 5’ terminus of the COI gene was amplified using standard primers (LepF1–LepR1) following established protocols (Smith et al. 2006, 2007, 2008). If the initial 658 bp amplification was not successful, composite sequences were generated using internal primers. Primer information for individual sequences can be retrieved from the Barcode of Life Data System (BOLD) (Ratnasingham and Hebert 2007), but primers are as detailed in Smith et al. (2008). DNA barcoding data and related information for all specimens studied in this paper can be accessed at: https://doi.org/10.5883/DATASET-ACGAP1.

Sequences were considered as “barcode-compliant” when they had 500 or more base pairs and correctly placed the species with its conspecifics in an NJ tree, the function of which is simple identification (as well as species discovery). We did not follow the more formal, but less pragmatically useful, definition of also having less than 1% ambiguous characters (Barcode Compliance standards as in http://www.boldsystems.org/index.php/resources/handbook?chapter=6_managingdata.html&section=record_list), though in selecting the holotype we strive to choose a specimen with a “full” and “complete” DNA barcode. The number of sequences available in BOLD for each species that we treat is provided in the “Molecular data” section within the taxonomic treatment of every species and all sequences can be obtained from BOLD.

In the taxonomic treatment of species, “Material Examined” presents the specimen’s information in the following format: “Number of females/males, acronym of the storing institution between parenthesis, COUNTRY: State/Province, city, other locality details, coordinates (in Decimal Degrees), date, collector name, biological information on host (starting with “ex”), ACG database codes (in the format “yy-SRNP-xxxxxx” for the host or “DHJPARxxxxxxx” for the wasp)”. For states of the United States and for Canadian provinces/territories, acronyms consisting of two capital letters are used, following Canada Post (http://www.canadapost.ca/tools/pg/manual/PGaddress-e.asp).

For phylogenetic analyses conducted outside of BOLD, DNA barcode sequences from 180 Mesoamerican Apanteles species were downloaded from BOLD along with 29 sequences from species of other Microgastrinae genera (Microplitis, Pholetesor, Parapanteles, Dolichogenidea). All sequences were imported into Geneious Pro 6.1 (Drummond et al. 2011) and aligned using the default settings for MUSCLE. The first and last nucleotide positions of the aligned dataset were deleted to reduce the amount of missing data. All sequences downloaded were over 500 bp long and the final aligned dataset contained 657 characters of which 315 were parsimony informative. The dataset was partitioned into two partitions, the first containing first and second codon positions and the second containing third codon positions. Model testing done in JModelTest v0.1.1 (Posada 2008) using the Bayesian Information Criterion selected the GTR+I+G model for both partitions. Two independent Bayesian analyses with 4 chains each were run in MrBayes v3.2.1 (Ronquist and Huelsenbeck 2003) for 31 253 000 generations each. Trace files of all parameters were examined in Tracer v1.5 (Rambaut and Drummond 2009) to verify that the runs had converged on the same stationary distribution, and to select the percentage of samples to remove as burn-in. A 10% burn-in was removed from both tree files which were then combined and resampled at 10%. A Maximum Clade Credibility Tree (MCCT) was made from the combined resampled tree file in TreeAnotator v1.7.5 (Rambaut and Drummond 2013). The MCCT only shows posterior probability values over 0.50 (Fig. 1). When discussing the support to particular clades (species-groups), those values are mentioned within the text in the format: “PP: 0.95” (i.e., posterior probability of 0.95). While this tree is obviously better than a simple NJ tree for displaying presumed phylogenetic relationships, we stress that the purpose of this paper is alpha taxonomy and not phylogeny, and any phylogenetic analyses are highly preliminary until hundreds more of the ACG and Mesoamerican species of Apanteles can be included.

Figure 1.

Bayesian Maximum Clade Credibility Tree (MCCT) for 180 species of Mesoamerican Apanteles with over 500 bp in the barcoding region and 29 species from other genera used as outgroups. Posterior probabilities over 0.50 are shown on the left side of nodes. Scale bar indicates branch length, expressed as the expected number of substitutions per site.

A neighbor-joining (Saitou and Nei 1987) tree was also constructed in Geneious Pro 6.1 (Drummond et al. 2011) using the TN93 model (Tamura and Nei 1993) (Fig. 2).

Figure 2.

Neighbor-joining Tree for 180 species of Mesoamerican Apanteles with over 500 bp in the barcoding region and 29 species from other genera used as outgroups. Scale bar indicates branch length, expressed as the expected number of substitutions per site.

Results
Definition of the genus Apanteles sensu stricto

The limits of Apanteles and other genera of Microgastrinae have lately been the subject of debate (e.g., van Achterberg 2003), and some recent references use a rather broad generic concept, e.g., Fauna Europaea (van Achterberg 2012), and Taxapad (Yu et al. 2012). We follow here the generic concepts proposed by Mason (1981), which have been widely adopted in parts of Europe (Papp 1988, Broad et al. 2012), Australia (Austin and Dangerfield 1992), China (Chen and Song 2004), and the New World (Whitfield 1995 and 1997, Fernández-Triana 2010, Fernández-Triana et al. 2013). For the purpose of this review, we consider the genera Choeras, Dolichogenidea, Exoryza, Iconella, and Illidops to be valid and not synonyms or subgenera of Apanteles sensu stricto. We provide below definitions of all of these genera, and our reasons for keeping them separate from Apanteles sensu stricto.

Choeras was defined by Mason (1981) to include species with a median carina on the propodeum (the Greek name of the genus precisely refers to that character). In addition, many species, although not all, have a more or less complete areolet (i.e. second submarginal cell) on the fore wing. The genus is far from being resolved, as Mason himself stated in the original description. Based on extensive material worldwide that we have been able to study, some species groups (e.g. the psarae group, as defined by Nixon (1965) and followed by Mason (1981), and many undescribed species) currently thought to be Choeras may be better placed in a different (new) genus; this applies especially to most of the species from the Oriental and Australasian regions. However, all of those species have a more or less complete median carina on the propodeum, and never have any indication of a propodeal areola (in contrast to Apanteles sensu stricto). DNA barcoding tends to clearly cluster the species of both genera separately (e.g. Smith et al. 2013).

Dolichogenidea, described by Viereck (1911), is perhaps the closest genus to Apanteles sensu stricto, and also the most controversial and difficult to separate from Apanteles. Mason (1981: 53–54) devoted several pages to discussing the main defining characters and the difficulties in separating both genera; he concluded, rather hopelessly, that in practice the decision was still somewhat arbitrary (because of the apparent continuum of variation in character). We have examined thousands of specimens from the world fauna of Microgastrinae, and have found that the only reliable character is the number and density of setae fringing on the median portion of the vannal lobe. Dolichogenidea has a convex to almost straight vannal lobe, which is uniformly fringed by setae. In Apanteles the vannal lobe is strongly concave to almost straight, and is lacking setae at midlength; the lack of setae may be partial (i.e. there may be some small and sparse setae on the lobe) or total (i.e. no setae at all). The differences between vannal lobes of those two genera were illustrated by Whitfield (1997: 364, figures 92–94). Both Mason (1981) and Whitfield (1997) discussed other characters that work in some (but not all) cases. Apart from morphology, DNA barcoding tends to clearly cluster the species of both genera separately (e.g. Fernández-Triana 2010; Smith et al. 2013). Some differences in host ranges and geographical distribution have also been observed, but no comprehensive revision of the data is available yet.

The genus Exoryza was described by Mason (1981) to include two species. It was considered to be mainly distributed in temperate regions, although two additional species recently described, one from the Neotropics (Valerio et al. 2004) and one from China (Song and Chen 2003) suggest that the genus is more widely distributed. Valerio et al. (2004) considered it to be closely related to both Dolichogenidea and Apanteles (and in a lesser extent also to Parapanteles and Pholetesor). While its generic position is still unclear, the presence of a convex, straight vannal lobe, uniformly fringed by setae (similar to that of Dolichogenidea) is the main feature that distinguishes this genus from Apanteles.

Mason (1981) described Iconella as a new genus based on the sinuous vein cu-a in the hind wing as a plesiomorphic character that suggests its unique status among similar genera. Besides that, Fernández-Triana et al. (2013) also considered the presence of a median longitudinal carina on the propodeum (or the secondary loss of that carina, which occurs in some Palaeartic species but not in the New World species) as a strong support for its generic status. DNA barcoding tends to clearly cluster the species of both Iconella and Apanteles separately (e.g. Fernández-Triana et al. 2013; Smith et al. 2013).

Mason (1981) erected Illidops to accommodate a group of species with the lower margin of the eyes converging and metasomal terga 3–7 weakly sclerotized. However, those characters are not universal within the genus, being absent in several species. This might be one reason why the genus has not been universally recognized. After studying species from different regions of the world, we found features that permit better definition of the genus Illidops, such as a band of rugosity centrally on the posterior edge of the scutellar disc; a shortened fore wing vein R1; and propodeum fully sculptured but without areola (instead, with a series of short carinae medially on the posterior 0.2–0.3 of the propodeum near the nucha). DNA barcoding tends to clearly cluster species of Illidops and Apanteles separately (e.g. Smith et al. 2013).

Summarizing, our definition of Apanteles sensu stricto, as used in this study is based on the following characters: a) propodeum never with a median longitudinal carina; either with carinae defining a partial or complete areola (sometimes areola obscured by propodeum sculpture, but still evident as an impression) or, very rarely, without any definition of an areola; b) Vannal lobe margin strongly concave to almost straight, and lacking setae at midlength (if some setae are present, they are small and sparsely distributed); c) posterior edge of the scutellar disc smooth; d) vein cu-a in the hind wing not sinuate. There are other useful characters to distinguish the genus (e.g. see Mason 1981, Whitfield 1997); but the four detailed above serve well to separate the Apanteles sensu stricto described here from Choeras (character a), from Dolichogenidea (character b), from Exoryza (character b), from Iconella (character d), and from Illidops (character c).

A total of 23 Apanteles species were previously known from Mesoamerica (Yu et al. 2012, Smith et al. 2013). After studying the type material for these taxa, 19 are retained in Apanteles sensu stricto and treated in this paper, while four species are transferred to other Microgastrinae genera. The status of the Mesoamerican species Apanteles albinervis (Cameron 1904) is revised, and one European species that is involved in a secondary homonym with the former is transferred to Iconella. All of these nomenclatorial acts are detailed below.

When the 186 new species from ACG described in this paper are included, the Apanteles fauna of Mesoamerica comprises 205 species, or about 10 times more species than previously known from that region. There was only one described species recorded from Costa Rica, 19 from Mesoamerica, 86 from the Neotropical region, and 1010 worldwide (Yu et al. 2012).

This study emphasizes how much is still unknown about the diversity of parasitoid wasps in general, and Microgastrinae in particular (e.g., Rodriguez et al. 2012). It is unlikely that ACG contains 20% of the species of a global genus as widely distributed and diverse as Apanteles. A more logical explanation is that, whenever other regions are as comprehensively studied, many more undescribed species of Apanteles will be revealed.

Even for ACG we are far from completing the inventory of Apanteles. We are aware of another 19 species (which would represent an additional 10% of increase for ACG), which we had to exclude because the available specimens were in poor condition and/or were only represented by males (in most cases Microgastrinae male specimens cannot be taxonomically dealt with, except through their DNA barcodes, or by inference through membership in a presumed sib group containing females, or reared from only one caterpillar). Those species are not described in this paper, although some of their interim names are provided here, for future reference, Apanteles Janzen11, Apanteles Janzen16, Apanteles Janzen34, Apanteles Rodriguez50, Apanteles Rodriguez74, Apanteles Rodriguez75, Apanteles Rodriguez79, Apanteles Rodriguez109, Apanteles Rodriguez121, Apanteles Rodriguez127, Apanteles Rodriguez128, Apanteles Rodriguez138, Apanteles Rodriguez143, Apanteles Rodriguez149, Apanteles Rodriguez161, Apanteles Rodriguez185, Apanteles Rodriguez200, Apanteles Rodriguez216, and Apanteles Rodriguez250. Full details of these presumed species and many more can be found in the ACG database online (http://janzen.sas.upenn.edu/caterpillars/database.lasso). We will describe these species in subsequent papers on the ACG Microgastrinae.

Species formerly described as Apanteles but here excluded from the genus
1. Dolichogenidea hedyleptae (Muesebeck, 1958), comb. n.
Apanteles hedyleptae Muesebeck, 1958: 443. (Puerto Rico, Trinidad and Tobago).
Note.

After examinating the holotype (NMNH), we consider this to be a species of Dolichogenidea-based on the evenly convex and uniformly setose vannal lobe.

2. Dolichogenidea politiventris (Muesebeck, 1958), comb. n.
Apanteles politiventris Muesebeck, 1958: 436. (Puerto Rico).
Note.

After examinating the holotype (NMNH), we consider this to be a species of Dolichogenidea-based on the evenly convex and uniformly setose vannal lobe.

3. Iconella albinervis (Tobias, 1964), stat rev.
Apanteles albinervis Tobias, 1964: 221. (Kazakhstan). Secondary homonym of Apanteles albinervis (Cameron, 1904).
Apanteles albinervis Tobias, 1964. Kept as a valid species by Shenefelt (1972: 438) as a result of an invalid replacement name for Apanteles albinervis (Cameron, 1904).
Iconella albinervis (Tobias, 1964). Transferred by Papp (1988: 151).
Apanteles albinervis Tobias, 1964. Transferred by van Achterberg (2003: 27).
Remarks.

The name Apanteles albinervis Tobias, 1964 could be considered a secondary homonym of Apanteles albinervis (Cameron, 1904) [see more explanations below, under the taxonomic treatment of Apanteles albinervis (Cameron, 1904) in the section “Taxonomic treatment of the Apanteles species in Mesoamerica, alphabetically by species-groups”]. However, Papp (1988) transferred the Tobias’ species to Iconella when treating the Palaearctic species under the (then new) reclassification of Microgastrinae proposed by Mason (1981). In a later paper, van Achterberg (2003) rejected Papp’s work and sank several genera of Microgastrinae under Apanteles, with Iconella being one of them. As part of those changes, Apanteles albinervis Tobias, 1964 was reinstated as a valid name – and, if accepted at present, would become a secondary homonym of Apanteles albinervis (Cameron, 1904). However, van Achterberg’s proposal is far from being widely accepted (Broad et al. 2012; Fernández-Triana 2010). We think that the best approach at present is to consider Iconella as a valid genus – not a junior synonym of Apanteles. Thus, we here transfer Tobias’s species back to Iconella, as done earlier by Papp (1988) – an arrangement also accepted by other workers (e.g., Inanç 1997, Kotenko 2007).

Article 59.2 of the International Code of Zoological Nomenclature (ICZN 1999) regulates the case of “Secondary homonyms not replaced when no longer considered congeneric” and states that “If in a case of secondary homonymy the junior species-group name has not been replaced [Art. 60], and the relevant taxa are no longer considered congeneric, the junior name is not to be rejected, even if one species-group name was originally proposed in the current genus of the other”. In accordance with that, the junior name Iconella albinervis (Tobias, 1964) is no longer considered congeneric with the senior name Apanteles albinervis (Cameron, 1904) and thus there is no need for a replacement name for the Tobias species. However, given previous changes in the generic status of this species, it should be kept in mind that, if future studies bring it back into Apanteles, at that moment a replacement name will be needed to avoid a secondary homonym with Apanteles albinervis (Cameron, 1904).

4. Illidops scutellaris (Muesebeck, 1921), comb. rev.
Apanteles scutellaris Muesebeck, 1921: 533.
Illidops scutellaris (Muesebeck). Papp 1988: 150.
Apanteles scutellaris Muesebeck. Austin and Dangerfield 1992: 9; Whitfield 1995: 247; van Achterberg 2003: 29.
Remarks.

After examining the holotype (NMNH), we consider this species as belonging to Illidops, as pointed out a quarter of a century ago by Papp (1988). Papp transferred the species from Apanteles to Illidops in his treatment of the European fauna, when he adopted the then recent reclassification of Microgastrinae by Mason (1981). Strangely, his action was overlooked, ignored or rejected by the taxonomic community, and even authors following the Mason system of genera (e.g., Austin and Dangerfield 1992, Whitfield 1995) did not treat the species as Illidops but kept it as Apanteles. Van Achterberg (2003) considered Illidops as a subgenus of Apanteles, and transferred all of the European species, including Illidops scutellaris, back to Apanteles. All of the character defining Illidops (as outlined in the above section “Definition of the genus Apanteles sensu stricto”) are present in Illidops scutellaris, and thus we here transfer the species Apanteles scutellaris back to Illidops.

5. Rhygoplitis sanctivincenti (Ashmead, 1900), comb. n.
Apanteles sanctivincenti Ashmead, 1900: 279. (Saint Vincent).
Remarks.

The species Apanteles sanctivincenti Ashmead, 1900 was described from a single male, but the type has never been found in the BMNH and is probably lost (Gavin Broad, personal communication). Thus, later researchers reviewing the genus (Muesebeck 1921, Nixon 1965) or cataloguing it (Szépligeti 1904, Shenefelt 1972) were unable to study it, and could only rely upon the very poor original description and key from Ashmead (1900: 279–280). Those five papers are the only publications citing the name Apanteles sanctivincenti Ashmead, and the species has been considered as belonging to Apanteles since its original description. However, after Mason’s (1981) paper splitting Apanteles into several genera, it is evident that Apanteles sanctivincenti Ashmead belongs to a different genus, based on its pronotum with a median longitudinal carina, a character that immediately excludes it from the current limits of Apanteles, but that occurs in several other genera of Microgastrinae. In his paper revising the fauna of the Caribbean islands of St. Vincent and Grenada, Ashmead (1900) treated five other genera of microgastrines: Microplitis, Protapanteles, Protomicroplitis, Urogaster and Pseudapanteles. The first three belong to completely different groups which can safely be excluded from the present analysis. Urogaster is no longer a valid genus (the majority of its species have been transferred to Apanteles). Pseudapanteles can also be excluded because its species have a median longitudinal groove on the first mediotergite, a trait not present in Apanteles sanctivincenti Ashmead, according to the original description. After carefully considering the distribution of other genera in the region, and comparing it with other species descriptions from the same paper (Ashmead 1900), we believe that the best generic placement for this species is Rhygoplitis.

It is worth mentioning that Ashmead (1900: 291) described two other species, Urogaster aciculatus and Pseudapanteles sancti-vincentis, which are now considered to be the same and to belong to Rhygoplitis; the valid species name currently is Rhygoplitis aciculatus. It is possible that Apanteles sanctivincenti is yet another name for that same species, meaning that three different names in three different genera were applied to the same species by the same author in the same paper! This case is not unlikely, due to Ashmead’s poor knowledge of the Microgastrinae (Mason 1981). In fact, the descriptions in his 1900 paper are not only very inconsistent (characters in the key do not correspond well to the descriptions, descriptions are not homogeneous, some body areas are named differently in the same paper, e.g., knees and femur) but they are also misleading, e.g., the original description of Urogaster aciculatus mentions the propodeum with a large, round areola, when it actually has no areola at all. We studied the three descriptions in detail to see if they could correspond to the same species. The lack of uniformity and different terminology prevents a certain conclusion, but they are similar in many regards, differing in minor details such as coloration (which may be meaningless anyway, because of the very small number of specimens examined by the author). Because the holotype of Apanteles sanctivincenti is lost, this situation may never be resolved unambiguously. Thus for the sake of name stability, and pending future studies on the genus, we just transfer Apanteles sanctivincenti to Rhygoplitis.

ACG species wrongly assigned to Apanteles in the past

In a first, non-taxonomic analysis of the Microgastrinae fauna of ACG, Smith et al. (2008) included 136 interim, unnamed, species of Apanteles – detailed in their “Datasets 1 and 2” of their “Supporting Information”. After reviewing those specimens, we feel that 12 of those species are better placed in other microgastrine genera and therefore transfer them here (Table 1). The rest are described below, together with additional ACG species identified after the publication of the Smith et al. (2008) paper.

Table 1.

Species considered as belonging to Apanteles by Smith et al. (2008) but transferred to other genera of Microgastrinae in the present paper. After the new assigned genus we provide the interim specific name whenever available (the format being “Genus Interim name”, e.g., Dolichogenidea Janzen90). The interim names allow for contemporary retrieval of full information of specimens in the online ACG database (http://janzen.sas.upenn.edu/caterpillars/database.lasso) as well as BOLD (www.barcodinglife.org). When those species are revised and published in their respective generic revisions, they will receive an appropriate formal scientific name.

Species name in Smith et al. (2008) New generic and/or interim species name assigned here
Apanteles Rodriguez02 Parapanteles Rodriguez02
Apanteles Rodriguez45 Parapanteles Whitfield45
Apanteles Rodriguez90 Dolichogenidea Janzen90
Apanteles Rodriguez102 Parapanteles Whitfield102
Apanteles Rodriguez118 Glyptapanteles Whitfield175
Apanteles Rodriguez119 Dolichogenidea Janzen119
Apanteles Rodriguez133 Parapanteles Whitfield133
Apanteles Rodriguez136 Parapanteles Whitfield302
Apanteles Rodriguez137 Parapanteles Whitfield303
Apanteles Rodriguez157 The name was applied to several specimens that might represent more than one species. The ones we could study are Dolichogenidea, the others are left as Apanteles Rodriguez157 and will be dealt with in future papers.
Apanteles Rodriguez164 Probably Dolichogenidea.
Apanteles Rodriguez172 Probably Dolichogenidea. We have only seen one specimen lacking legs, antenna and metasoma, which cannot be properly assigned to genus until more material is available.
General comments on the biology and morphology of Apanteles in Mesoamerica

At present we have biological information (host records, solitary/gregariousness of wasp larvae) for 169 (82%) of the described species of Apanteles in Mesoamerica. Some records may be questionable, especially early citations of hosts for Apanteles carpatus (Say, 1836) which may be incorrect (this is a cosmopolitan species and examination of vouchers from all biogeographic regions is needed to solve the problem). But for the vast majority of species (especially all of the ones reared in ACG) the records are accurate, – and comprehensive enough to draw conclusions on host relationships at a higher taxon level.

Twenty Lepidoptera families have been recorded as hosts of Apanteles in Mesoamerica (or 14 families, if six only recorded from Apanteles carpatus, – which are likely to be wrong –, are excluded). Most hosts species belong to just three families: Hesperiidae (33%), Elachistidae (26%) and Crambidae (21%), distantly followed by Pyralidae (4%), Choreutidae (3%) and Gelechiidae (3%) (Fig. 3). However, the boundary between ACG Elachistidae and Gelechiidae is very poorly defined. When the moth taxa are clearly worked out, the ratio of these two families may be quite different.

Figure 3.

Proportion of Lepidoptera families parasitized by 169 species of Apanteles with known host records in Mesoamerica (data source mainly from the ACG inventory).

Most of the ACG reared Apanteles species (154 species, or 91%) are monophagous or oligophagous, attacking one host family, and usually only one or a very few species within the same genus. Only 15 species (9%) are somewhat polyphagous, parasitizing hosts in two or more Lepidoptera families, but even they tend to be very selective in their hosts. Furthermore, larger sample sizes and better DNA barcode data has generally shown that these “somewhat polyphagous” species in ACG are often complexes of narrow specialists, as for example the case of what was believed to be “Apanteles leucostigmus” (Smith et al. 2008).

Outside ACG, there is not enough data to assess if narrow host ranges per microgastrine wasp species is a widespread phenomenon (although unpublished evidence suggest that this might be the case and there is no reason to think that ACG Apanteles are abnormal). If this pattern proves to be commonplace worldwide, it will have a strong influence on biological control and on biodiversity studies (e.g., Rodriguez et al. 2012, Smith et al. 2013).

A total of 98 species (58%) of the Mesoamerican Apanteles with associated biological information have multiple larvae developing in one host caterpillar (gregarious), and these are generally viewed as originating from a single ovipositing female. In contrast, 71 species (42%) are solitary, having just one larva per parasitized caterpillar. Although there is no comparable information from other regions, about 80% of the Nearctic species of Apanteles with reliable data available are solitary (Whitfield unpublished data).

Most wasp cocoons (either solitary or gregarious) are stuck to the leaf substrate, and over, under, or near the host cadaver, which “lives” only a few days, if at all, after emergence of the wasp larvae. As cocoon structure often appears a species-level characteristic, it is shown for each species when possible (Figs 210330).

The fauna of Mesoamerica, especially that of ACG, seems to have some peculiar morphological characteristics. For example, one species in ACG is the only known Apanteles in the world with partially white genae. That feature is present in Alphomelon and occasionally in a few other genera of Microgastrinae (e.g., Mason 1981, Deans et al. 2003), but had never before been found in Apanteles.

Although orange-yellow coloration is not uncommon in tropical Apanteles, it is mostly restricted to legs, portions of metasoma, and, rarely, spots on the mesosoma. Four ACG species (2%) are the first known members of the genus to have extensive orange coloration, including the whole head. Interestingly, none of these four species seem to be closely related.

Similarly, only five ACG species have pectinate tarsal claws, while one species has cleft tarsal claws. The vast majority (97.5%) of the Mesoamerican species either have simple tarsal claws, or with 1–2 basal spine-like setae.

About 10% of the Mesoamerican Apanteles within several groups (including anabellecordobae, which is the third largest species-group in the region) have the hypopygium either unfolded or with only 1–3 pleats. That is very unusual in Apanteles and may force a future redefinition of Apanteles limits.

Almost one quarter of the Apanteles species in Mesoamerica have a somewhat elongate glossa, although it is never as large and bilobate as in some other characteristic genera of Microgastrinae such as Pseudapanteles, Promicrogaster, etc.

Species groups of Mesoamerican Apanteles

In order to deal with its high diversity, the genus Apanteles has been partitioned into species groups since 1880. Mason (1981) provides a summary of current understanding of the evolution of those groups as well as references to different papers on the topic. A total of 44 species-groups for the world fauna were proposed by Nixon (1965), an arrangement that has generally been accepted and incorporated into subsequent revisions, e.g., Mason (1981) and European fauna (revised by Papp between 1976 and 1990).

While some of these species groups appear to represent monophyletic or at least morphologically coherent groups, many are poorly defined, and some are just containers for species that do not fit into any other group. To further complicate things, many species have never been assigned to a particular species-group (e.g., only half of the previously described species of Mesoamerican Apanteles had been assigned to a group before this paper).

In spite of the shortcomings in the species-group system, it remains a useful tool for partitioning the large number of Apanteles species. Until a more comprehensive, phylogeny-based taxonomy is available, groups of species based on inference from morphology remain the most practical approach.

For the Mesoamerican region we recognize and propose 32 species-groups of Apanteles (Table 2) and we assign most of the species known for the region to one of them. All groups are new, except for two (Apanteles ater and Apanteles diatraeae) previously created and used by several authors (e.g., Nixon 1965, Mason 1981, Austin and Dangerfield 1989, Whitfield et al. 2001, 2002). For 30 species we did not have strong support to assign them to any of the 32 established groups; and neither the morphological, molecular nor biological data are sufficient to justify them as individual groups. Those 30 species are left out of groups, and categorized as “unassigned”.

Table 2.

Species-groups of Apanteles in Mesoamerica, in alphabetical order. # of Spp: Total number of species currently within a group. Lepidoptera host families: “?” Unknown; for the carpatus group “??” refers to a mix of old and questionable references including nine different families (Gelechiidae, Lasiocampidae, Lecithoceridae, Lymantriidae, Pyralidae, Thaumetopoeidae, Tineidae, Tortricidae, Zygaenidae). Larvae: S–wasp larvae solitary; S?–wasp larvae strongly suspected to be solitary but not conclusive evidence; G–wasp larvae gregarious. ?–Unknown. In cases where a species-group has both solitary and gregarious larvae, the most common occurrence is indicated first. MOR, DNA, BIO: degree of group support by morphological (MOR), molecular (DNA), and biological (BIO) data. “+” Strong support, “-”No support, “P” Partial support, “?” Unknown.

Species-group # of Spp Lepidoptera host families Larvae MOR DNA B IO
adelinamoralesae 19 Elachistidae, Pyralidae G, S + + +
adrianachavarriae 9 Attevidae, Crambidae, Elachistidae, Tortricididae S, G P P P
adrianaguilarae 3 Tortricidae G + + +
alejandromorai 13 Elachistidae, Gelechiidae S + + +
anabellecordobae 14 Hesperiidae G, S + + +
anamarencoae 2 Elachistidae, Oecophoridae, Tortricidae S + - -
arielopezi 2 Elachistidae, Tortricidae G + + ?
ater 9 Crambidae, Pyralidae S - - ?
bernyapui 4 Crambidae, Elachistidae, Gelechiidae, Noctuidae S + + ?
bienvenidachavarriae 3 Elachistidae S + P +
calixtomoragai 3 Hesperiidae S + + +
carlosguadamuzi 6 Choreutidae, Crambidae, Gelechiidae, Elachistidae G, S + + ?
carlosrodriguezi 3 Elachistidae, Choreutidae, Crambidae G, S + P P
carloszunigai 2 ? ? + + ?
carpatus 5 ?? G + + ?
coffeellae 4 Gracillariidae, Lyonetiidae S? P - ?
diatraeae 3 Crambidae G + + +
dickyui 2 ? ? + + ?
erickduartei 5 Crambidae S + + +
glenriverai 2 Pyralidae G + + +
guadaluperodriguezae 2 Crambidae G + - +
humbertolopezi 2 Elachistidae S + - ?
isidrochaconi 2 ? ? + + ?
javierobandoi 2 Choreutidae S + - ?
Joserasi 2 Hesperiidae S + + +
keineraragoni 2 Crambidae, Riodinidae G, S + - -
Leucostigmus 39 Hesperiidae G + + +
marisolnavarroae 2 Pyralidae S + + +
megathymi 2 Hesperiidae G + ? -
paranthrenidis 4 Pyralidae, Crambidae, Gelechiidae, Noctuidae, Sesiidae S, G + ? -
ronaldgutierrezi 2 Choreutidae S + P ?
Samarshalli 2 ? ? + + ?

Six groups each have nine or more species, jointly representing half of all described Mesoamerican species. The largest are the leucostigmus group (39 species parasitizing Hesperiidae) and the adelinamoralesae group (19 species attacking Elachistidae), both with many more Mesoamerican species awaiting description.

A total of 15 species-groups (47%) are represented by two species. This situation is mainly due to the fact that only the ACG fauna has been comprehensively studied. When the remainder of the Mesoamerican Apanteles fauna is revised, many of these groups are likely to have more species associated. We have seen in collections numerous undescribed species from the Neotropics other than ACG, species that fall into some of the new groups erected here (to encourage further study in the future we have noted those cases in the taxonomy treatment of species below).

Most groups (79%) are strongly supported by at least two of three sets of independent data: molecular (DNA barcodes), biology (host), and morphology (Table 2), and their component species can be clearly recognized and delimited. Others are defined mainly by shared morphological characters. In one case (Apanteles joserasi), the group could only be defined by a unique combination of hosts and barcoding characters (see couplet 29 of the key to species-groups).

Some groups partially overlap with others, e.g., the adrianachavarriae, javierobandoi, and joserasi groups, and future studies may reveal that they should be merged. However, without study of the whole Mesoamerican (or even Neotropical) fauna, we preferred to take a conservative approach in accommodating the perceived differences.

The non-ACG ater, coffeellae, megathymi, and paranthrenidis groups could not be defined unambiguously, and should only be considered as interim groupings of species; they will need to be revisited when more studies on the world fauna are undertaken.

The species aidalopezae and leonelgarayi (currently not assigned to any group), and the groups carlosrodriguezi and samarshalli, all comprise species that might be better placed in other genera in the future. For example, the samarshalli group clusters out of all other Apanteles species, strongly indicating (PP: 1.0 in the Bayesian analysis, Fig. 1) that its two species may best be placed in a (new) different genus. However, pending a comprehensive phylogenetic study of Microgastrinae, we decided that it is best to here describe all those species as belonging to Apanteles. The key to species groups separate those species in the first four couplets.

Key to the species-groups of Mesoamerican Apanteles

[This section provides a key to all species-groups of Apanteles in Mesoamerica, including 30 species that could not be assigned to any current group and are keyed individually throughout the key. It is followed by keys to species within every species-group (the groups arranged in alphabetical order). After all keys, standardized descriptions of every species are provided (the species arranged in alphabetical order). To facilitate finding individual species, Table 3 provides alphabetical lists of species and species-groups].

1 Fore wing with vein 2M very short, its anterior half very close to anterior half of vein 2RS, in a way that obliterates most of space of second submarginal cell (Figs 160b, 205a, c); antenna very short, 0.5 × body length, and not surpassing posterior margin of mesosoma (Figs 160e, 205a–c), and body not distinctly flattened dorsoventrally, and pro- and meso- femora yellow, and pterostigma relatively broad, its length less than 2.7 × its width [Distribution: Canada (ON), Costa Rica (ACG), Mexico and US (FL)] samarshalli species-group [2 species]
Fore wing with vein 2M completely separated from vein 2RS (as in Fig. 4b); antenna usually as long or longer than body length, at least surpassing posterior margin of mesosoma; if antenna shorter (i.e., not surpassing posterior margin of mesosoma), then body distinctly flattened dorsoventrally (as in Fig. 203a), and/or profemur (partially or entirely) and mesofemur dark brown to black, and/or pterostigma usually relatively narrow, its length more than 3.0 × its width 2
2(1) Ovipositor sheaths extremely short, 0.3 × or less metatibia length (Figs 138a, c); T2 relatively large, its median length 0.7–0.9 × as long as T3 median length (Fig. 138f); T1 mostly smooth (except for 2–3 small carinae centrally); body with extensive yellow-orange coloration (all legs except for metatarsus and posterior 0.2 of metatibia, tegula and humeral complex, all laterotergites and sternites, hypopygium) Apanteles leonelgarayi Fernández-Triana, sp. n.
Ovipositor sheaths at least 0.4 × as long as metatibia (usually much more than that); T2 median length much shorter than T3 median length (almost always 0.5 × or less); T1 almost always with some sculpture; body color variable 3
3(2) Hypopygium with a relatively wide but short fold, restricted to posterior 0.4–0.5 of hypopygium length, where no pleats are visible (or, rarely, at most with a single, weakly marked pleat); ovipositor short and slightly to strongly curved downwards (Figs 36a, c); ovipositor sheaths very short (0.4–0.5 × as long as metatibia, Fig. 36c); relatively small size, body length and fore wing length not surpassing 2.5 mm 4
Hypopygium usually with large fold and numerous pleats, if rarely with no visible pleats or just one pleat, then ovipositor relatively long and thick, not strongly curved downwards, and/or ovipositor sheaths longer than 0.5 × metatibia length (usually much longer), and/or body length and fore wing length surpassing 2.5 mm 5
4(3) Pterostigma white (Fig. 36b); glossa elongate; antenna much shorter than body, not extending beyond mesosoma (Fig. 36a) Apanteles aidalopezae Fernández-Triana, sp. n.
Pterostigma brown, with small pale spot at base (Fig. 96b); glossa not elongate; antenna usually as long as body or slightly shorter (extending beyond mesosoma) carlosrodriguezi species-group [3 species]
5(3) Head entirely orange (except for black interocellar area and/or small spot on upper part of gena), anteromesoscutum, scutellar disc, and axillar complex completely or almost completely orange (Figs 37, 135, 139, 163) 6
Head mostly black to dark brown (except for clypeus and labrum, which may be yellow-orange) or head black with gena partially white; anteromesoscutum and scutellar disc usually black to dark brown, at most with relatively small yellow or orange spots 9
6(5) Mesopleuron and mesosternum dark brown to black, except for upper anterior and/or lower posterior corners of mesopleuron which are orange (Figs 37a, 163a) 7
Mesopleuron either completely orange, or mostly orange (upper anterior 1/3 dark brown to black), mesosternum fully orange (Figs 135a, 139a) 8
7(6) Mesoscutellar disc smooth (Fig. 163g); all mediotergites dark brown to black (Fig. 163g); tarsal claws pectinate Apanteles waldymedinai Fernández-Triana, sp. n.
Mesoscutellar disc mostly punctured (Fig. 37e); T1 mostly orange and T3 partially yellow (Fig. 37e); tarsal claws with one basal spine-like seta Apanteles alejandromasisi Fernández-Triana, sp. n.
8(6) T1 mostly white except for small black spot posteriorly (Fig. 135f); all laterotergites, most sternites, and hypopygium white; scutoscutellar sulcus almost obliterated, with less than 4 small impressions (Fig. 135f); propodeal areola open basally and without transverse carinae; tarsal claws pectinate Apanteles juliodiazi Fernández-Triana, sp. n.
Metasoma entirely black (Fig. 139f); scutoscutellar sulcus not obliterated, with 5–6 small impressions (Fig. 139f); propodeal areola closed basally and with transverse carinae extending to spiracle; tarsal claws with one basal spine-like seta Apanteles luisgaritai Fernández-Triana, sp. n.
9(5) Head with gena partially white (Figs 155a, d) Apanteles rogerblancoi Fernández-Triana, sp. n.
Head with gena entirely black 10
10(9) Tarsal claws pectinate, or cleft with a basal spine [Hosts: Hesperiidae, Pyrginae] 11
Tarsal claws either simple or with 1–2 basal spine-like setae 12
11(10) T1 coarsely sculptured with longitudinal and transverse striation; T2 with some sculpture, especially along posterior margin (Fig. 119f); tarsal claws cleft and with a basal spine Apanteles garygibsoni Fernández-Triana, sp. n.
T1 smooth, at most with fine sculpture along lateral margins; T2 smooth and polished (Fig. 87e); tarsal claws pectinate calixtomoragai species-group [3 species]
12(10) Smaller individuals, body length 1.6–2.2 mm, and fore wing length 1.8–2.4 mm; and body distinctly flattened (as in Figs 203a, 204a) or T1 length >3.5 × its posterior width (as in Figs 106g, 107f, 108f, 203g) 13
Larger individuals, body length and fore wing lengths usually more than 2.5 mm; if rarely less than 2.5 mm, then body not distinctly flattened and T1 length <3.5 × its posterior width 17
13(12) All legs, including coxae (except for small spot on anterior 0.2 of metacoxa), entirely yellow or whitish-yellow (Figs 83a, c, g) Apanteles bettymarchenae Fernández-Triana, sp. n.
At least metacoxa completely, and part of femora and tibiae, with dark brown to black coloration 14
14(13) Antenna as long as or longer than body length; T1 strongly narrowing toward apex from its apical half — ratio of T1 basal width/T1 apical width >2.0 × and ratio of T1 length/T1 apical width >3.5 × (Fig. 106g); body not distinctly flattened; parasites of leaf-mining Lepidoptera. [Hosts: Gracillariidae, Lyonetiidae. Distribution: Costa Rica (ACG), Guadeloupe, Puerto Rico] coffeellae species-group [4 species]
Antenna shorter, its length at most 0.7 × body length, usually much less; T1 usually narrowing toward apex less strongly — ratio of T1 basal width/T1 apical width <2.0 ×, ratio of T1 length/T1 apical width usually <3.5 × (Figs 203g, 204g); body distinctly flattened dorsoventrally (as in Figs 203a, 204a); parasites of non-mining Lepidoptera 15
15(14) Smooth area on lateral face of scutellum very narrow and small, its maximum height at most 0.2 × lateral face height (Fig. 67e); hypopygium inflexible, without any fold [Hosts: Tortricidae] Apanteles anapiedrae Fernández-Triana, sp. n.
Smooth area on lateral face of scutellum at least 0.4 × lateral face height (usually much more) (Fig. 122f, 203g, 204g); hypopygium with a translucid median fold with at least one pleat visible 16
16(15) Propodeal areola open anteriorly, elongate and more or less parallel-sided (Figs 203g, 204g), its maximum width (at around half of propodeum length) <1.3 × its width at posterior end (nucha); hypopygium with a wide median fold with usually four or more visible pleats [Hosts: stem-boring Crambidae] diatraeae species-group [3 species]
Propodeal areola clearly closed anteriorly and widening centrally, its maximum width (at around half of propodeum length) >1.5 × its width at apex (nucha); hypopygium with a translucid median fold with 1–3 visible pleats [Hosts: leaf-folder Crambidae] guadaluperodriguezae species-group [2 species]
17(12) Hypopygium with outer margin inflexible, without a median fold (as in Figs 51c, 56c), or hypopygium with a median, transparent, semi-desclerotized fold with none or very few (usually 1–3) pleats occupying just outermost area of fold (as in Figs 52c, 55c) 18
Hypopygium with outer margin with a wide median, transparent, semi-desclerotized fold, with 4 or more pleats occupying most or whole fold (as in Fig. 145d) 23
18(17) Ovipositor relatively thick, as thick or thicker than width of median flagellomerus, and with basal width 3.0–5.0 × its apical width posterior to constriction (Figs 51c, 52a, c, 54c, 56c) anabellecordobae species-group [14 species]
Ovipositor relatively thin, thinner than width of median flagellomerus, and with basal width <2.0 × its apical width after constriction (as in Figs 68a, c, 142c) 19
19(18) T1 mostly smooth (Fig. 156g); T1 slightly widening from anterior margin to 0.7–0.8 × mediotergite length (where maximum width is reached), then narrowing towards posterior margin Apanteles rolandovegai Fernández-Triana, sp. n.
T1 mostly sculptured, at least on posterior half (Figs 68g, 142f); T1 more or less parallel-sided for its entire length, or parallel-sided for 0.5–0.7 × its length then narrowing posteriorly so mediotergite anterior width >1.1 × posterior width (Figs 68g, 142f) 20
20(19) T1 length 1.4 × its width; fore wing length 3.3 mm Apanteles marialuisariasae Fernández-Triana, sp. n.
T1 length at least 2.3 × its width; fore wing length at most 2.8 mm 21
21(20) All coxae, profemur partially, and meso- and metafemora completely, dark brown to black (Fig. 68a); mesoscutellar disc mostly smooth (Fig. 68g); hypopygium with outer margin inflexible, without a median fold Apanteles andreacalvoae Fernández-Triana, sp. n.
At least pro- and mesocoxae (and usually metacoxa), pro- and mesofemora, and most of metafemur (except for apical 0.2 or less), yellow to orange (Figs 99a, c, 149a, c); mesoscutellar disc mostly punctured, or with punctures near margins and centrally smooth (Figs 99g, 149f); hypopygium with a median, transparent, semi-desclerotized fold with none or very few (usually 1–3) pleats occupying just outermost area of fold 22
22(20) Flagellomerus 14 1.0 × as long as wide; scutoscutellar sulcus with 9 impressed pits; tarsal claws with one basal spine-like seta; T1 length 2.3 × its width; T2 with some sculpture near its posterior margin (Fig. 149f) Apanteles oscarchavesi Fernández-Triana, sp. n.
Flagellomerus 14 at least 1.6 × as long as wide; scutoscutellar sulcus with 5–6 impressed pits; tarsal claws simple; T1 length at least 3.2 × its width; T2 mostly smooth (Fig. 99g) carloszunigai species-group [2 species]
23(17) T2 broadly rectangular, its apical width 2.2 × or less than its median length (as in Figs 38e, 39g, 40f, 105g, 112f) 24
T2 transverse and relatively narrow, its apical width 2.5 × or more its median length 26
24(23) Ovipositor relatively thick (Fig. 112c), as thick or thicker than width of median flagellomerus, and with basal width 3.0–5.0 × its apical width posterior to constriction [Hosts: Hesperiidae. Distribution: ACG] Apanteles diegotorresi Fernández-Triana, sp. n.
Ovipositor relatively thin (as in Fig. 38a), thinner than width of median flagellomerus, and with basal width <2.0 × its apical width after constriction [Hosts: Elachistidae. Distribution: ACG] 25
25(24) Ovipositor sheaths more than 1.2 × as long as metatibia, and usually longer than metasoma (as in Fig. 38a); fore wing with maximum width of first discal cell at most 1.1 × its maximum height (usually 1.0 × or less), second abscissa of vein 1CU slightly curved (as in Figs 38b, 39b, 40b, 41b, 42b, 43b, 44b, 46b); T1 less than 3.3 × as long as its apical width alejandromorai species-group [13 species]
Ovipositor sheaths less than 1.0 × as long as metatibia, and much shorter than metasoma (Fig. 105a); fore wing with maximum width of first discal cell 1.3 × its maximum height, second abscissa of vein 1CU straight (Fig. 105b); T1 more than 3.4 × as long as its apical width Apanteles christianzunigai Fernández-Triana, sp. n.
26(23) Pterostigma relatively broad, its length less than 3.0 × its width (as in Fig. 104b), and T2 mostly sculptured with strong longitudinal striation (Figs 102g, 103g, 104g) carpatus species-group [5 species]
Pterostigma relatively narrow, its length more than 3.0 × its width, and T2 either smooth or weakly sculptured, without strong longitudinal striation 27
27(26) Ovipositor relatively thick and strong, as thick or thicker than width of median flagellomerus and with basal width 3.0-5.0 × its apical width posterior to constriction (Figs 133a, c, 168c, 172c, 179c) 28
Ovipositor relatively thin, thinner than width of median flagellomerus, and with basal width <2.0 × its apical width after constriction 30
28(27) Maximum height of mesoscutellum lunules 0.4 × maximum height of lateral face of mesoscutellum (Fig. 120f); antenna shorter than body length; propodeum usually evenly sculptured in most of its surface (Fig. 120f) [Hosts: Pyralidae] glenriverai species-group [2 species]
Maximum height of mesoscutellum lunules 0.7 × or more maximum height of lateral face of mesoscutellum (as in Fig. 133f); antenna as long or longer than body length; propodeum with strong sculpture limited to anterior half, posterior half mostly smooth and shiny; propodeum with transverse carinae complete and strongly raised, clearly delimited from background sculpture (as in Fig. 133f) [Hosts: Hesperiidae] 29
29(28) Solitary parasitoids of Venada (Hesperiidae); cocoons as in Fig. 291 [See comments under species-group treatment for further justification on its status] joserasi species-group [2 species, one undescribed]
Gregarious parasitoids of several genera of Hesperiidae but not Venada; cocoons as in Figs 304329 leucostigmus species-group [39 species]
30(27) Body with extensive yellow and/or orange coloration, including tegula and humeral complex, parts of the axillar complex, sometimes posterior margin of mesoscutum (right in front of scutoscutellar sulcus), all coxae (rarely metacoxa dark brown to black), sometimes lateral edges of T3 and T4, most of laterotergites 1–4, most sternites and hypopygium (partial or completely) (as in Figs 33a, f, 114a, f, 127a, f, 141a, f, 159g, 161a, c) 31
Body with much less extensive yellow-orange coloration: usually metacoxa (and sometimes also pro- and meso- coxae) partially to completely reddish, brown or black; axillar complex, tergites, most of laterotergites, and hypopygium (partial or completely) dark brown to black; tegula and humeral complex color variable but rarely both yellow 43
31(30) T2 mostly sculptured (Fig. 159g) Apanteles rosibelelizondoae Fernández-Triana, sp. n.
T2 mostly smooth, at most with some sculpture near the posterior margin 32
32(30) T1 length at least 3.8 × (usually more than 4.0 ×) its width at posterior margin (Fig. 141f, 161h) and ovipositor sheaths 0.4 × as long as metatibia (Fig. 161a, c) 33
T1 length at most 3.2 × its width at posterior margin and/or ovipositor sheaths at least 0.6 × as long as metatibia 34
33(32) Body length 3.0–3.2 mm, forewing length 3.1–3.3 mm; tegula and humeral complex dark brown; anteromesoscutum with two orange spots laterally near posterior margin (Fig. 141f); tarsal claws simple; ocular-ocellar line 2.1 × as long as posterior ocellus diameter; interocellar distance 1.6 × posterior ocellus diameter; flagellomerus 14 1.2 × as long as wide; flagellomerus 2 2.5 × as long as flagellomerus 14; T2 width at posterior margin 3.0–3.4 × its medial length [Hosts: Crambidae] Apanteles marcogonzalezi Fernández-Triana, sp. n.
Body length 2.4–2.7 mm, forewing length 2.6–2.7 mm; tegula and humeral complex yellow; anteromesoscutum entirely black (Fig. 161h); tarsal claws with one basal spine-like seta; ocular-ocellar line 2.5 × as long as posterior ocellus diameter; interocellar distance 2.0 × posterior ocellus diameter; flagellomerus 14 1.6 × as long as wide; flagellomerus 2 2.0 × as long as flagellomerus 14; T2 width at posterior margin 3.6 × its medial length [Hosts: Choreutidae] Apanteles sergiocascantei Fernández-Triana, sp. n.
34(32) T1 length 1.8 × its width; and mesoscutellar disc smooth (Fig. 162e); and fore wing vein 2RS more than 2.0 × as long as vein 2M; and T2 width at posterior margin 3.9 × its length; and ocular-ocellar line 1.6 × posterior ocellus diameter [Hosts: Crambidae borers, Diatraea spp.] Apanteles vulgaris Fernández-Triana, sp. n.
T1 length usually more than 2.3 × its width; and/or mesoscutellar disc with punctures; and/or fore wing with vein 2RS less than 2.0 × as long as vein 2M; and/or T2 width at posterior margin less than 3.6 × its length; and/or ocular-ocellar line at least 1.7 × posterior ocellus diameter [Hosts: Choreutidae, Elachistidae, Gelechiidae, Tortricidae; if Crambidae, not borers] 35
35(34) Ovipositor sheaths 1.3 × as long as metatibia (Figs 127a, c, 128a, c); and body length and fore wing length 4.0 mm; and mesoscutellar disc smooth centrally (Figs 127f, 128f); and ocular-ocellar line 1.5 × posterior ocellus diameter isidrochaconi species-group [2 species]
Ovipositor sheaths usually shorter than metatibia (rarely 1.1–1.2 ×); body length and fore wing length usually less than 3.0 mm, if more than that (up to 3.5 mm) then mesocutellar disc punctured, and/or ocular-ocellar line at least 2.0 × posterior ocellus diameter 36
36(35) Metacoxae entirely or mostly (anterior 0.5 or more) dark brown to black (as in Figs 34a, 115f) 37
All coxae entirely white, yellow or bright orange, at most with small brown spot on anterior 0.1 or less (as in Figs 33f, 114f, 116a, f) 38
37(36) Fore wing with length of vein r 1.4 × or less length of vein 2RS; ocular-ocellar line at least 2.4 × posterior ocellus diameter; T2 width at posterior margin at least 3.7 × its length adrianaguilarae species-group (in part) [3 species]
Fore wing with length of vein r 2.4 × length of vein 2RS; ocular-ocellar line 2.2 × posterior ocellus diameter; T2 width at posterior margin at least 3.3 × its length erickduartei species-group (in part) [5 species]
38(36) Ocular-ocellar line 2.5 × as long as posterior ocellus diameter; and T2 width at posterior margin at least 4.0 × (usually more) as long as its medial length; and fore wing with vein 2M as long as vein (RS+M)b adrianaguilarae species-group (in part) [3 species]
Ocular-ocellar line at most 2.2 × as long as posterior ocellus diameter; and/or T2 width at posterior margin usually 3.5 × (or much less) as long as its medial length; and/or fore wing with vein 2M usually shorter than vein (RS+M)b 39
39(38) Mesoscutellar disc mostly punctured (as in Figs 114f, 115f) 40
Mesoscutellar disc mostly smooth, at most with few, scattered punctures near margins, central part smooth (as in Figs 80f, 81g, 134f); if rarely mostly punctured, then posterior 0.2–0.3 of anteromesoscutum (especially centrally and along posterior margin) and upper anterior corner of mesopleura orange (as in Figs 80f, 82g) 41
40(39) Ovipositor sheaths clearly as long or longer as metatibia (1.0–1.2 ×, rarely 0.9 ×); tarsal claws with one basal spine-like seta erickduartei species-group (in part) [5 species]
Ovipositor sheaths clearly shorter than metatibia (0.4 ×) (Figs 118a, c); tarsal claws simple Apanteles flormoralesae Fernández-Triana, sp. n.
41(39) T1 mostly sculptured, with excavated area centrally with transverse striation inside and polished knob centrally on posterior margin of mediotergite and T1 mostly parallel–sided for 0.5–0.7 of its length, then narrowing posteriorly so mediotergite anterior width >1.1 × posterior width (Fig. 134f), and anteromesoscutum and T1 entirely black; T2 width at posterior margin 5.4 × its length; metafemur length 3.5 × its width Apanteles juanhernandezi Fernández-Triana, sp. n.
T1 mostly smooth (as in Fig. 90g), if mostly sculptured, then T1 mostly parallel-sided (as in Fig. 79g), or anteromesoscutum with posterior 0.2 orange (as in Fig. 80f) and/or T1 orange to light-brown; T2 width at posterior margin at most 4.0 × (usually much less) its length; metafemur length at most 3.2 × its width (usually 3.0 × or less) 42
42(41) T1 almost always black, same color of propodeum (some decoloured specimens may have T1 dark brown); T1 length at most 2.3 × its width, and mostly strongly sculptured, with longitudinal striation laterally and a central excavated area with transverse striation (Fig. 79g) bernyapui species-group [4 species]
T1 orange-yellow, orange or light brown, always lighter than propodeum color (as in Fig. 90g); T1 length at least 2.5 × its width (usually much more), with some weak sculpture on posterior 0.2–0.5 but mostly looking smooth (Fig. 90g) carlosguadamuzi species-group [6 species]
43(30) Tegula different in color from humeral complex 44
Tegula same color as humeral complex 57
44(43) Pterostigma mostly transparent or white, with thin brown borders; and all coxae dark brown to black 45
Pterostigma either fully brown, mostly brown (at most with small pale area centrally or anteriorly), or fully white, without brown borders; and/or procoxa (sometimes also mesocoxa) yellow-orange to light brown 51
45(44) T1 at most 1.3 × as long as wide at posterior margin, and T2 mostly smooth and/or pro- and mesocoxae yellow paranthrenidis species-group [4 species]
T1 at least 1.7 × as long as wide (usually much more), if rarely 1.3 ×, then T2 fully sculptured with longitudinal striation and all coxae dark brown to black 46
46(45) Glossa elongate in both sexes (Fig. 140e) [Hosts: Crambidae. Distribution: ACG] Apanteles luisvargasi Fernández-Triana, sp. n.
Glossa not elongate 47
47(46) Ovipositor sheaths usually more than 1.2 × metatibial length, if rarely 1.0–1.1 ×, then profemur at least partially, and meso- and metafemora completely, dark brown to black; and tegula yellow-white; and fore wing with vein 2RS less than 2.0 × length of vein 2M (usually less than 1.6 ×); and humeral complex half yellow-white, half dark brown; and T2 width at posterior margin at least 2.9 × its length [Host: Elachistidae] adelinamoralesae species-group [19 species]
Ovipositor sheaths usually less than 1.1 × metatibial length; if rarely more than 1.2 ×, then pro- and mesofemora completely, and metafemur at least partially, yellow-orange; and/or fore wing with vein 2RS more than 2.0 × length of vein 2M; and/or humeral complex usually unicolour (or very rarely humeral complex half yellow-white, half dark, and tegula dark brown); and/or T2 width at posterior margin at most 2.7 × its length 48
48(47) Propodeum with sculpture on anterior half different from posterior half (which is either smoother or with clearly different pattern of sculpture than the anterior half) (Figs 23e, 24e, 25f, 26f, 27f, 28f, 29f, 30g, 31g, 84g, 85f, 86f) 49
Propodeum fully sculptured, without much difference in sculpture between anterior and posterior halves (as in Fig. 146f) 50
49(48) T2 width at posterior margin at most 2.7 × (usually 2.5 × or less) its length (Figs 84g, 85f, 86f) [Hosts: Elachistidae] bienvenidachavarriae species-group [3 species]
T2 width at posterior margin at least 3.5 × its length (Figs 23e, 24e, 25f, 26f, 27f, 28f, 29f, 30g, 31g) [Hosts: Crambidae, Tortricidae, Yponomeutidae] adrianachavarriae species-group [9 species]
50(48) Profemur partially, and meso- and metafemora completely, dark brown to black; ovipositor relatively thick (anterior width 2.0 × as posterior width) (Fig. 132c) josediazi species-group
Pro- and mesofemora completely to partially yellow-orange; ovipositor relatively thin, about same width throughout its length (Fig. 146c) megathymi species-group [2 species]
51(44) Ovipositor sheaths 1.4–1.5 × as long as metatibia length (Figs 129a, c); and body length and fore wing length at least 3.2 mm Apanteles isidrovillegasi Fernández-Triana, sp. n.
Ovipositor sheaths usually less than 1.1 × as long as metatibia length; if rarely ovipositor sheaths 1.3 × as long as metatibia length, then body length and fore wing length at most 2.2 mm 52
52(51) Glossa elongate (Fig. 143e) [Host: Elachistidae] Apanteles mariamendezae Fernández-Triana, sp. n.
Glossa not elongate 53
53(51) T1 mostly parallel-sided for 0.7 of its length, then strongly narrowing posteriorly so T1 length at least 3.0 × its width at posterior margin (Fig. 124f) [Host: Riodinidae] Apanteles hectorsolisi Fernández-Triana, sp. n.
T1 either clearly widening towards posterior margin, or slightly widening from anterior margin to 0.7–0.8 mediotergite length (where maximum width is reached), then narrowing towards posterior margin, or parallel-sided so T1 length at most 2.5 × its width at posterior margin (usually much less than that) 54
54(53) T1 clearly widening towards posterior margin, 1.3 × as long as wide at posterior margin; T2 with posterior margin sinuate (Fig. 35e), width at expanded central area 1.7 × as large as width at lateral area; T2 4.0 × as long as wide at posterior margin; ovipositor relatively thick, basal width about twice apical width Apanteles aichagirardae Fernández-Triana, sp. n.
T1 parallel-sided or slightly widening from anterior margin to 0.7–0.8 mediotergite length (where maximum width is reached), then narrowing towards posterior margin, so T1 length at least 2.0 × its width at posterior margin; T2 with posterior margin straight, thus central and lateral areas of same length (as in Fig. 69g); T2 usually less than 4.0 × as long as wide at posterior margin; ovipositor about same width throughout its length 55
55(54) Ovipositor sheaths 0.6 × as long as metatibia length (Figs 69a, c); humeral complex dark; metatrochanter, metatrochantellus, and anterior 0.2–0.3 of metafemur yellow or yellow-white (Fig. 69c) ronaldgutierrezi species-group [2 species]
Ovipositor sheaths usually as long as or longer than metatibia length, if slightly shorter (0.9 ×) then antenna shorter than body (its length not surpassing half of metasoma); humeral complex half pale, half dark; metafemur, metatrochanter, and sometimes metatrochantellus dark brown to black 56
56(55) Body length at most 2.4 mm, and fore wing length at most 2.7 mm; mesofemur fully yellow; mesoscutellar disc mostly smooth (Figs 69, 70); ovipositor sheaths at least 1.3 × as long as metatibia length or antenna shorter than body (its length not surpassing half of metasoma) [Hosts: Crambidae, Elachistidae, Riodinidae, Tortricidae] arielopezi species-group [2 species]
Body length at least 3.3 mm (usually more), and fore wing length at least 3.3 mm (usually more); mesofemur anterior 0.5–0.8 dark brown (Figs 144a, 145a); mesoscutellar disc mostly punctured; ovipositor sheaths at most 1.1 × as long as metatibia length [Hosts: Pyralidae] marisolnavarroae species-group [2 species]
57(43) T1 length more than 4.5 × its posterior width (Fig. 154g); vannal lobe straight and fully setose (with slightly shorter and sparser setae centrally); metacoxa partially yellow and partially dark brown (Fig. 154a) Apanteles robertmontanoi Fernández-Triana, sp. n.
T1 length less than 3.7 × its posterior width; vannal lobe usually strongly concave, centrally without setae (or with very small, very sparse setae); metacoxa usually entirely dark brown to black 58
58(57) Ovipositor sheaths 0.5 × as long as metatibia (Fig. 136a, c); and relatively small size, body length 2.3 mm, and fore wing length 2.4 mm; and metatibial spurs at most 0.4 × as long as first segment of metatarsus (Fig. 136c) keineraragoni species-group [2 species]
Ovipositor sheaths at least 0.8 × as long as metatibia; and/or relatively larger size, body length and fore wing length at least 2.5 mm; and/or metatibial spurs at least 0.5 × as long as first segment of metatarsus 59
59(58) Glossa elongate (Fig. 130e, 131e); and tarsal claws simple javierobandoi species-group [2 species]
Glossa not elongate; tarsal claws usually with single basal spine-like seta 60
60(59) Pterostigma entirely brown or brown with pale spot at base 61
Pterostigma entirely transparent or mostly transparent with only thin brown borders 63
61(60) At least pro- and mesocoxae entirely pale (white-yellow, yellow, or orange) (Figs 72a, 73a, 74a, 75a, 76a, 78a) ater species-group (in part)
All coxae dark brown to black 62
62(61) Tegula, humeral complex, all femora and tibiae yellow (metafemur with small brown spot on posterior 0.2 × or less) (Figs 65a, d, e, 66, a); T2 mostly smooth (Fig. 66f); ovipositor sheaths at least 1.4 × as long as metatibia length (Figs 65a, c, 66a, c) anamarencoae species-group [2 species]
Tegula, humeral complex, meso- and metafemora dark, metatibia (partially), and usually mesotibia (partially) dark brown to black (Figs 125a, c, f, 126a, c, g); T2 fully sculptured with longitudinal striation (Figs 125f, 126g); ovipositor sheaths at most 1.3 × (usually less than 1.0 ×) as long as metatibia length (Figs 125c, 126c) humbertolopezi species-group [2 species]
63(60) Tegula and humeral complex dark brown to black, or pro-, meso-, and part of metacoxae yellow-orange ater species-group (in part)
Tegula and humeral complex yellow, and meso- and metacoxae (sometimes also procoxa) dark brown to black 64
64(63) T1 length 1.5 × its width (Fig. 148f); T2 mostly smooth (Fig. 148f); body length 3.2 mm, and fore wing length 3.7 mm Apanteles monicachavarriae Fernández-Triana, sp. n.
T1 length at least 2.4 × its width (Figs 110g, 11f); T2 sculptured, mostly near posterior margin (Figs 110g, 11f); body length 2.2–2.6 mm, and fore wing length 2.2–2.6 mm dickyui species-group
Table 3.

Alphabetical lists of Mesoamerican Apanteles: by species (columns 1 and 2) and by species-groups (columns 3 and 4).

Ordered by species Ordered by species-groups
Species Species-group Species-group Species
Apanteles adelinamoralesae adelinamoralesae adelinamoralesae Apanteles adelinamoralesae
Apanteles adrianachavarriae adrianachavarriae Apanteles carloscastilloi
Apanteles adrianaguilarae adrianaguilarae Apanteles didiguadamuzi
Apanteles adrianguadamuzi adrianachavarriae Apanteles edgarjimenezi
Apanteles aichagirardae Unassigned Apanteles gerardosandovali
Apanteles aidalopezae Unassigned Apanteles isaacbermudezi
Apanteles albanjimenezi carpatus Apanteles jorgecortesi
Apanteles albinervis leucostigmus Apanteles juanvictori
Apanteles alejandromasisi Unassigned Apanteles juniorlopezi
Apanteles alejandromorai alejandromorai Apanteles laurenmoralesae
Apanteles alvarougaldei leucostigmus Apanteles leninguadamuzi
Apanteles anabellecordobae anabellecordobae Apanteles luiscanalesi
Apanteles anamarencoae anamarencoae Apanteles luislopezi
Apanteles anamartinezae adrianachavarriae Apanteles manuelarayai
Apanteles anapiedrae Unassigned Apanteles paulaixcamparijae
Apanteles anariasae ater Apanteles ronaldmurilloi
Apanteles andreacalvoae Unassigned Apanteles wilbertharayai
Apanteles angelsolisi leucostigmus Apanteles yolandarojasae
Apanteles arielopezi arielopezi Apanteles zeneidabolanosae
Apanteles balthazari megathymi adrianachavarriae Apanteles adrianachavarriae
Apanteles bernardoespinozai leucostigmus Apanteles adrianguadamuzi
Apanteles bernyapui bernyapui Apanteles anamartinezae
Apanteles bettymarchenae Unassigned Apanteles felipechavarriai
Apanteles bienvenidachavarriae bienvenidachavarriae Apanteles irenecarrilloi
Apanteles calixtomoragai calixtomoragai Apanteles luiscantillanoi
Apanteles carloscastilloi adelinamoralesae Apanteles mariatorrentesae
Apanteles carlosguadamuzi carlosguadamuzi Apanteles ronaldquirosi
Apanteles carlosrodriguezi carlosrodriguezi Apanteles yilbertalvaradoi
Apanteles carlosviquezi leucostigmus adrianaguilarae Apanteles adrianaguilarae
Apanteles carloszunigai carloszunigai Apanteles ivonnetranae
Apanteles carolinacanoae anabellecordobae Apanteles vannesabrenesae
Apanteles carpatus carpatus alejandromorai Apanteles alejandromorai
Apanteles christianzunigai Unassigned Apanteles deifiliadavilae
Apanteles cinthiabarrantesae carlosguadamuzi Apanteles eulogiosequeirai
Apanteles ciriloumanai leucostigmus Apanteles fernandochavarriai
Apanteles coffeellae coffeellae Apanteles franciscoramirezi
Apanteles cristianalemani ater Apanteles freddysalazari
Apanteles cynthiacorderoae leucostigmus Apanteles gabrielagutierrezae
Apanteles deifiliadavilae alejandromorai Apanteles juancarrilloi
Apanteles deplanatus diatraeae Apanteles luisbrizuelai
Apanteles diatraeae diatraeae Apanteles luisgarciai
Apanteles dickyui dickyui Apanteles marvinmendozai
Apanteles didiguadamuzi adelinamoralesae Apanteles minornavarroi
Apanteles diegoalpizari ater Apanteles tiboshartae
Apanteles diegotorresi Unassigned anabellecordobae Apanteles anabellecordobae
Apanteles diniamartinezae leucostigmus Apanteles carolinacanoae
Apanteles duniagarciae anabellecordobae Apanteles duniagarciae
Apanteles duvalierbricenoi leucostigmus Apanteles edwinapui
Apanteles edgarjimenezi adelinamoralesae Apanteles eldarayae
Apanteles edithlopezae carlosguadamuzi Apanteles freddyquesadai
Apanteles eduardoramirezi dickyui Apanteles guillermopereirai
Apanteles edwinapui anabellecordobae Apanteles harryramirezi
Apanteles eldarayae anabellecordobae Apanteles joseperezi
Apanteles eliethcantillanoae leucostigmus Apanteles luciariosae
Apanteles erickduartei erickduartei Apanteles manuelpereirai
Apanteles esthercentenoae paranthrenidis Apanteles marianopereirai
Apanteles eugeniaphilipsae leucostigmus Apanteles osvaldoespinozai
Apanteles eulogiosequeirai alejandromorai Apanteles ruthfrancoae
Apanteles federicomatarritai leucostigmus anamarencoae Apanteles anamarencoae
Apanteles felipechavarriai adrianachavarriae Apanteles juanlopezi
Apanteles felixcarmonai erickduartei arielopezi Apanteles arielopezi
Apanteles fernandochavarriai alejandromorai Apanteles mauriciogurdiani
Apanteles flormoralesae Unassigned ater Apanteles anariasae
Apanteles franciscopizarroi ater Apanteles cristianalemani
Apanteles franciscoramirezi alejandromorai Apanteles diegoalpizari
Apanteles freddyquesadai anabellecordobae Apanteles franciscopizarroi
Apanteles freddysalazari alejandromorai Apanteles galleriae
Apanteles fredi diatraeae Apanteles impiger
Apanteles gabrielagutierrezae alejandromorai Apanteles jairomoyai
Apanteles galleriae ater Apanteles josejaramilloi
Apanteles garygibsoni Unassigned Apanteles leucopus
Apanteles gerardobandoi leucostigmus bernyapui Apanteles bernyapui
Apanteles gerardosandovali adelinamoralesae Apanteles javiersihezari
Apanteles gladysrojasae leucostigmus Apanteles raulacevedoi
Apanteles glenriverai glenriverai Apanteles victorbarrantesi
Apanteles gloriasihezarae carlosrodriguezi bienvenidachavarriae Apanteles bienvenidachavarriae
Apanteles guadaluperodriguezae guadaluperodriguezae Apanteles josecalvoi
Apanteles guillermopereirai anabellecordobae Apanteles marisolarroyoae
Apanteles harryramirezi anabellecordobae calixtomoragai Apanteles calixtomoragai
Apanteles hazelcambroneroae leucostigmus Apanteles manuelriosi
Apanteles hectorsolisi Unassigned Apanteles petronariosae
Apanteles humbertolopezi humbertolopezi carlosguadamuzi Apanteles carlosguadamuzi
Apanteles impiger ater Apanteles cinthiabarrantesae
Apanteles inesolisae leucostigmus Apanteles edithlopezae
Apanteles insularis ronaldgutierrezi Apanteles javiercontrerasi
Apanteles irenecarrilloi adrianachavarriae Apanteles jesusbrenesi
Apanteles isaacbermudezi adelinamoralesae Apanteles williamcamposi
Apanteles isidrochaconi isidrochaconi carlosrodriguezi Apanteles carlosrodriguezi
Apanteles isidrovillegasi Unassigned Apanteles gloriasihezarae
Apanteles ivonnetranae adrianaguilarae Apanteles robertoespinozai
Apanteles jairomoyai ater carloszunigai Apanteles carloszunigai
Apanteles javiercontrerasi carlosguadamuzi Apanteles yeissonchavesi
Apanteles javierobandoi javierobandoi carpatus Apanteles albanjimenezi
Apanteles javiersihezari bernyapui Apanteles carpatus
Apanteles jesusbrenesi carlosguadamuzi Apanteles rhomboidalis
Apanteles jesusugaldei leucostigmus Apanteles robertovargasi
Apanteles jimmychevezi samarshalli Apanteles rolandoramosi
Apanteles johanvargasi leucostigmus coffeellae Apanteles coffeellae
Apanteles jorgecortesi adelinamoralesae Apanteles laurahuberae
Apanteles jorgehernandezi leucostigmus Apanteles lisabearssae
Apanteles josecalvoi bienvenidachavarriae Apanteles mariaguevarae
Apanteles josecortesi leucostigmus diatraeae Apanteles deplanatus
Apanteles josediazi Unassigned Apanteles diatraeae
Apanteles josejaramilloi ater Apanteles fredi
Apanteles josemonteroi leucostigmus dickyui Apanteles dickyui
Apanteles joseperezi anabellecordobae Apanteles eduardoramirezi
Apanteles joserasi joserasi erickduartei Apanteles erickduartei
Apanteles juanapui isidrochaconi Apanteles felixcarmonai
Apanteles juancarrilloi alejandromorai Apanteles luishernandezi
Apanteles juangazoi javierobandoi Apanteles milenagutierrezae
Apanteles juanhernandezi Unassigned Apanteles ronaldcastroi
Apanteles juanlopezi anamarencoae glenriverai Apanteles glenriverai
Apanteles juanmatai leucostigmus Apanteles pablovasquezi
Apanteles juanvictori adelinamoralesae guadaluperodriguezae Apanteles guadaluperodriguezae
Apanteles juliodiazi Unassigned Apanteles marcobustosi
Apanteles juniorlopezi adelinamoralesae humbertolopezi Apanteles humbertolopezi
Apanteles keineraragoni keineraragoni Apanteles pablotranai
Apanteles laurahuberae coffeellae isidrochaconi Apanteles isidrochaconi
Apanteles laurenmoralesae adelinamoralesae Apanteles juanapui
Apanteles leninguadamuzi adelinamoralesae javierobandoi Apanteles javierobandoi
Apanteles leonelgarayi Unassigned Apanteles juangazoi
Apanteles leucopus ater joserasi Apanteles joserasi
Apanteles leucostigmus leucostigmus keineraragoni Apanteles keineraragoni
Apanteles lilliammenae leucostigmus Apanteles ronaldnavarroi
Apanteles lisabearssae coffeellae leucostigmus Apanteles albinervis
Apanteles luciariosae anabellecordobae Apanteles alvarougaldei
Apanteles luisbrizuelai alejandromorai Apanteles angelsolisi
Apanteles luiscanalesi adelinamoralesae Apanteles bernardoespinozai
Apanteles luiscantillanoi adrianachavarriae Apanteles carlosviquezi
Apanteles luisgarciai alejandromorai Apanteles ciriloumanai
Apanteles luisgaritai Unassigned Apanteles cynthiacorderoae
Apanteles luishernandezi erickduartei Apanteles diniamartinezae
Apanteles luislopezi adelinamoralesae Apanteles duvalierbricenoi
Apanteles luisvargasi Unassigned Apanteles eliethcantillanoae
Apanteles luzmariaromeroae leucostigmus Apanteles eugeniaphilipsae
Apanteles manuelarayai adelinamoralesae Apanteles federicomatarritai
Apanteles manuelpereirai anabellecordobae Apanteles gerardobandoi
Apanteles manuelriosi calixtomoragai Apanteles gladysrojasae
Apanteles manuelzumbadoi leucostigmus Apanteles hazelcambroneroae
Apanteles marcobustosi guadaluperodriguezae Apanteles inesolisae
Apanteles marcogonzalezi Unassigned Apanteles jesusugaldei
Apanteles marcovenicioi leucostigmus Apanteles johanvargasi
Apanteles mariachavarriae leucostigmus Apanteles jorgehernandezi
Apanteles mariaguevarae coffeellae Apanteles josecortesi
Apanteles marialuisariasae Unassigned Apanteles josemonteroi
Apanteles mariamendezae Unassigned Apanteles juanmatai
Apanteles marianopereirai anabellecordobae Apanteles leucostigmus
Apanteles mariatorrentesae adrianachavarriae Apanteles lilliammenae
Apanteles marisolarroyoae bienvenidachavarriae Apanteles luzmariaromeroae
Apanteles marisolnavarroae marisolnavarroae Apanteles manuelzumbadoi
Apanteles marvinmendozai alejandromorai Apanteles marcovenicioi
Apanteles mauriciogurdiani arielopezi Apanteles mariachavarriae
Apanteles megastidis paranthrenidis Apanteles minorcarmonai
Apanteles megathymi megathymi Apanteles pabloumanai
Apanteles milenagutierrezae erickduartei Apanteles randallgarciai
Apanteles minorcarmonai leucostigmus Apanteles raulsolorsanoi
Apanteles minornavarroi alejandromorai Apanteles ricardocaleroi
Apanteles monicachavarriae Unassigned Apanteles rodrigogamezi
Apanteles oscarchavezi Unassigned Apanteles ronaldzunigai
Apanteles osvaldoespinozai anabellecordobae Apanteles rostermoragai
Apanteles pablotranai humbertolopezi Apanteles sergioriosi
Apanteles pabloumanai leucostigmus Apanteles sigifredomarini
Apanteles pablovasquezi glenriverai Apanteles wadyobandoi
Apanteles paranthrenidis paranthrenidis marisolnavarroae Apanteles marisolnavarroae
Apanteles paulaixcamparijae adelinamoralesae Apanteles randallmartinezi
Apanteles petronariosae calixtomoragai megathymi Apanteles balthazari
Apanteles randallgarciai leucostigmus Apanteles megathymi
Apanteles randallmartinezi marisolnavarroae paranthrenidis Apanteles esthercentenoae
Apanteles raulacevedoi bernyapui Apanteles megastidis
Apanteles raulsolorsanoi leucostigmus Apanteles paranthrenidis
Apanteles rhomboidalis carpatus Apanteles thurberiae
Apanteles ricardocaleroi leucostigmus ronaldgutierrezi Apanteles insularis
Apanteles robertmontanoi Unassigned Apanteles ronaldgutierrezi
Apanteles robertoespinozai carlosrodriguezi samarshalli Apanteles jimmychevezi
Apanteles robertovargasi carpatus Apanteles samarshalli
Apanteles rodrigogamezi leucostigmus Unassigned Apanteles aichagirardae
Apanteles rogerblancoi Unassigned Unassigned Apanteles aidalopezae
Apanteles rolandoramosi carpatus Unassigned Apanteles alejandromasisi
Apanteles rolandovegai Unassigned Unassigned Apanteles anapiedrae
Apanteles ronaldcastroi erickduartei Unassigned Apanteles andreacalvoae
Apanteles ronaldgutierrezi ronaldgutierrezi Unassigned Apanteles bettymarchenae
Apanteles ronaldmurilloi adelinamoralesae Unassigned Apanteles christianzunigai
Apanteles ronaldnavarroi keineraragoni Unassigned Apanteles diegotorresi
Apanteles ronaldquirosi adrianachavarriae Unassigned Apanteles flormoralesae
Apanteles ronaldzunigai leucostigmus Unassigned Apanteles garygibsoni
Apanteles rosibelelizondoae Unassigned Unassigned Apanteles hectorsolisi
Apanteles rostermoragai leucostigmus Unassigned Apanteles isidrovillegasi
Apanteles ruthfrancoae anabellecordobae Unassigned Apanteles josediazi
Apanteles samarshalli samarshalli Unassigned Apanteles juanhernandezi
Apanteles sergiocascantei Unassigned Unassigned Apanteles juliodiazi
Apanteles sergioriosi leucostigmus Unassigned Apanteles leonelgarayi
Apanteles sigifredomarini leucostigmus Unassigned Apanteles luisgaritai
Apanteles thurberiae paranthrenidis Unassigned Apanteles luisvargasi
Apanteles tiboshartae alejandromorai Unassigned Apanteles marcogonzalezi
Apanteles vannesabrenesae adrianaguilarae Unassigned Apanteles marialuisariasae
Apanteles victorbarrantesi bernyapui Unassigned Apanteles mariamendezae
Apanteles vulgaris Unassigned Unassigned Apanteles monicachavarriae
Apanteles wadyobandoi leucostigmus Unassigned Apanteles oscarchavezi
Apanteles waldymedinai Unassigned Unassigned Apanteles robertmontanoi
Apanteles wilbertharayai adelinamoralesae Unassigned Apanteles rogerblancoi
Apanteles williamcamposi carlosguadamuzi Unassigned Apanteles rolandovegai
Apanteles yeissonchavesi carloszunigai Unassigned Apanteles rosibelelizondoae
Apanteles yilbertalvaradoi adrianachavarriae Unassigned Apanteles sergiocascantei
Apanteles yolandarojasae adelinamoralesae Unassigned Apanteles vulgaris
Apanteles zeneidabolanosae adelinamoralesae Unassigned Apanteles waldymedinai
adelinamoralesae species-group

This group comprises 19 species, defined by having ovipositor sheaths usually >1.2 × metatibia length; femora mostly (except for posterior half of profemur) dark brown to black; tegula yellow-white and humeral complex half yellow-white, half dark brown; and mediotergite 2 width at posterior margin at least 2.9 × its median length. The group is supported by the Bayesian molecular analysis (PP: 0.5 for the whole group, most of its species have PP between 0.9–1.0; Fig. 1). Hosts: Elachistidae and on two occasions, Pyralidae. All described are from ACG, but many species attacking elachistids in Mesoamerica are likely to be part of this group.

Key to species of the adelinamoralesae group
1 Metatibia entirely or mostly (>0.7 posteriorly) dark brown to black, with yellow-orange coloration restricted to anterior 0.2 or less (as in Figs 4a, 6c, 12c, a, 14a) 2
Metatibia yellow-orange at least on anterior 0.5 (usually more), with dark brown to black coloration restricted to posterior 0.5 or less (as in Figs 7c, 9a, c, 18c) 11
2(1) Ovipositor sheaths 1.0–1.1 × as long as metatibia 3
Ovipositor sheaths 1.3–1.6 × as long as metatibia 5
3(2) T1 parallel-sided for 0.7–0.8 of its length, then narrowing posteriorly so mediotergite anterior width >1.1 × posterior width; T2 width at posterior margin 4.4 × its medial length (Fig. 21h); metafemur 2.7 × as long as wide (Fig. 21c) Apanteles yolandarojasae Fernández-Triana, sp. n.
T1 slightly widening from anterior margin to 0.7–0.8 mediotergite length (where maximum width is reached), then narrowing towards posterior margin, with widest part of tergite (centrally) being 1.2 × that of base and/or apex; T2 width at posterior margin at most 3.1 × its medial length (as in Fig. 12f); metafemur at least 2.9 × as long as wide (Figs 12c, 17c) 4
4(3) Flagellomerus 2 2.4 × as long as wide; flagellomerus 14 1.3 × as long as wide; metafemur 3.3 × as long as wide; fore wing with vein 2RS 1.9 × as long as vein 2M Apanteles juniorlopezi Fernández-Triana, sp. n.
Flagellomerus 2 2.9 × as long as wide; flagellomerus 14 1.7 × as long as wide; metafemur 2.9 × as long as wide; fore wing with vein 2RS 1.1 × as long as vein 2M Apanteles manuelarayai Fernández-Triana, sp. n. (N = 5)
5(2) Mesoscutellar disc mostly smooth (Figs 4e, 22g); tarsal claws simple 6
Mesoscutellar disc mostly punctured, or at least with punctures near margins; tarsal claws with single basal spine-like seta 7
6(5) Metatibia with inner spur 2.0 × as long as outer spur; flagellomerus 2 2.2 × as long as wide; T1 2.0 × as long as wide at posterior margin; fore wing with vein r 1.2 × as long as vein 2RS, and vein 2RS 1.5 × as long as vein 2M [Cocoons: Gregarious. Hosts: Lethata trochalosticta] Apanteles zeneidabolanosae Fernández-Triana, sp. n.
Metatibia with inner spur 1.3 × as long as outer spur; flagellomerus 2 2.9 × as long as wide; T1 2.6 × as long as wide at posterior margin; fore wing with vein r 1.8 × as long as vein 2RS, and vein 2RS 2.1 × as long as vein 2M [Cocoons: Solitary. Hosts: Antaeotricha sp., Stenoma sp.] Apanteles adelinamoralesae Fernández-Triana, sp. n. (N = 1)
7(5) Fore wing with vein r at most 1.4 × as long as vein 2RS; metafemur 3.4 × as long as wide; interocellar distance usually 1.5 × as long as ocellus diameter (rarely up to 1.7 ×); ovipositor sheaths at least 1.6 × as long as metatibia (very rarely 1.5 ×) (Fig. 6c); protibia completely yellow (Fig. 6a) Apanteles didiguadamuzi Fernández-Triana, sp. n.
Fore wing with vein r at least 1.6 × as long as vein 2RS (usually more than 1.7 ×); metafemur at most 3.3 × as long as wide (usually much less); interocellar distance usually more than 1.8 × as long as ocellus diameter (rarely 1.7 ×); ovipositor sheaths at most 1.4 × as long as metatibia, usually less (very rarely 1.5 ×) (Figs 8a, 11a, 14c, 16a, c); protibia with anterior 0.5 yellow, posterior 0.5 dark brown to black (As in Fig. 16a) 8
8(7) T2 width at posterior margin at most 2.9 × its length (Fig. 8f); metafemur length 3.3 × its width Apanteles gerardosandovali Fernández-Triana, sp. n. (N = 5)
T2 width at posterior margin at least 3.2 × its length (usually 3.5 ×) (Figs 11f, 14f, 16f); metafemur length at most 3.0 × its width 9
9(8) T2 mostly smooth; T2 width at posterior margin at most 3.2 × its length (Figs 11f) Apanteles juanvictori Fernández-Triana, sp. n.
T2 usually with some sculpture; T2 width at posterior margin at least 3.5 × its length (Figs 14f, 16f) 10
10(9) Fore wing with vein r 2.3 × as long as vein 2RS, and vein 2RS 1.2 × as long as vein 2M; mesoscutellar disc with punctures near margins, central part mostly smooth [Hosts: Anadasmus sp., Cerconota sp.] Apanteles luislopezi Fernández-Triana, sp. n.
Fore wing with vein r 1.8 × as long as vein 2RS, and vein 2RS 1.5 × as long as vein 2M; mesoscutellar disc mostly punctured [Hosts: Stenoma byssina] Apanteles leninguadamuzi Fernández-Triana, sp. n.
11(1) Ovipositor sheaths 1.0 × as long as metatibia (rarely 1.1 ×) (Fig. 9c) Apanteles isaacbermudezi Fernández-Triana, sp. n. (N = 4)
Ovipositor sheaths 1.3-1.6 × as long as metatibia (rarely 1.2 ×) (Figs 5a, 7a, 10a, 13a) 12
12(11) T2 width at posterior margin at most 2.9 × its length, if rarely 3.0–3.2 × then T1 length at least 2.1 × its width at posterior margin and fore wing vein 2RS as long as vein 2M and vein 2M as long as vein (RS+M)b 13
T2 width at posterior margin at least 3.2 × its length (usually much more), and/or T1 length less than 2.0 × its width at posterior margin and/or fore wing vein 2RS longer than vein 2M and/or vein 2M shorter than vein (RS+M)b 15
13(12) Tarsal claws simple; fore wing with vein r 1.6 × as long as vein 2Rs, vein 2RS 1.6 × as long as vein 2M, and vein 2M 0.6 × as long as vein (RS+M)b Apanteles edgarjimenezi Fernández-Triana, sp. n.
Tarsal claws with single basal spine-like seta; fore wing with vein r at least 1.7 × as long as vein 2Rs, vein 2RS at most 1.3 × as long as vein 2M, and vein 2M at least 0.9 × as long as vein (RS+M)b 14
14(13) Interocellar distance at most 2.0 × ocellus diameter (usually less than 1.8 ×); mesoscutellar disc with punctures near the margin, central part mostly smooth; T1 length 2.1 × its width at posterior margin; ovipositor sheaths usually 1.5–1.6 × as long as metatibia length; if very rarely ovipositor sheaths 1.3 × as long as metatibia length, then body length and fore wing length 2.0 mm (otherwise body and fore wing length 2.9–3.3 mm) Apanteles carloscastilloi Fernández-Triana, sp. n.
Interocellar distance 2.1 × ocellus diameter; mesoscutellar disc mostly punctured; T1 length 1.7 × its width at posterior margin; ovipositor sheaths usually 1.3–1.4 × as long as metatibia length; body length 2.9–3.0 mm; fore wing length 3.1–3.4 mm Apanteles jorgecortesi Fernández-Triana, sp. n.
15(12) Ovipositor sheaths 1.6 × as long as metatibia; flagellomerus 2 2.5 × as long as wide; metatibial inner spur 1.7 × as long as outer spur Apanteles laurenmoralesae Fernández-Triana, sp. n.
Ovipositor sheaths at most 1.4 × as long as metatibia; flagellomerus 2 at least 2.6 × as long as wide (usually 2.9 × or more); metatibial inner spur at most 1.5 × as long as outer spur (usually less than 1.4 ×) 16
16(15) T2 fully sculptured; T2 width at posterior margin 4.6 × its length (Fig. 20g); body length 3.2 mm; fore wing length 3.4 mm Apanteles wilbertharayai Fernández-Triana, sp. n. (N = 1)
T2 mostly smooth, at most with weak and sparse punctures laterally near posterior margin; T2 width at posterior margin at most 3.6 × its length (Figs 15f, 18g, 19g); body length and fore wing length usually less than 3.0 mm (if rarely over 3.2 mm, then T2 width at posterior margin at most 3.2 × its length) 17
17(16) Interocellar distance 2.2 × as long as posterior ocellus diameter; mesoscutellar disc with punctures near the margin, central part mostly smooth Apanteles luiscanalesi Fernández-Triana, sp. n.
Interocellar distance 1.8 × as long as posterior ocellus diameter; mesoscutellar disc mostly punctured 18
18(17) T1 parallel-sided; T2 with some sculpture, mostly near posterior margin; fore wing with vein 2RS 1.0 × as long as vein 2M; outer margin of hypopygium extending about the same length of last tergites Apanteles paulaixcamparijae Fernández-Triana, sp. n.
T1 slightly widening from anterior margin to 0.7–0.8 mediotergite length (where maximum width is reached), then narrowing towards posterior margin; T2 mostly smooth; fore wing with vein 2RS 1.5 × as long as vein 2M; outer margin of hypopygium clearly extending beyond last tergites Apanteles ronaldmurilloi Fernández-Triana, sp. n.
adrianachavarriae species-group

This group comprises nine species with mesofemur, metafemur and all or most of metatibia dark brown to black; pterostigma with thin brown borders, centrally white or translucid; and mediotergite 1 with strong longitudinal striations. The group is likely to be artificial, at least partially, and it may end being part of a larger group (including the current joserasi javierobandoi groups). However, morphology, host data and DNA barcoding (Fig. 1), provide some support for most of its component species; and it seems better to keep this group separated for the time being. Hosts: Attevidae, Crambidae, Elachistidae, and Tortricidae. One of the species within this group, Apanteles felipechavarriai, is only known from a female in poor condition and cannot be keyed out using morphology alone beyond couplet 3 of the key below, thus barcoding data was used to distinguish that species from the remainder. All species described in this group are from ACG.

Key to species of the adrianachavarriae group
1 Metatibia with black coloration at most on posterior 0.3–0.5 (Figs 29a, c) [Hosts: Crambidae, Leucochromodes sp.] Apanteles mariatorrentesae Fernández-Triana, sp. n. (N = 2)
Metatibia almost completely black, except for anterior 0.2 or less which is yellow (as in Figs 23c, 25d, 27c, 28a, c, 30a, 31c) 2
2(1) T1 length at least 2.1 × its width at posterior margin and T2 width at posterior margin at most 4.0 × its length (if rarely T1 length 1.9 × its width at posterior margin, then T2 width at posterior margin less than 3.6 × its length) 3
T1 length at most 1.7 × (usually 1.6 × or less) its width at posterior margin and T2 width at posterior margin at least 4.3 × (usually 4.4 × or more) its length 7
3(2) A total of 18 diagnostic characters in the barcoding region: 81 G, 82 C, 99 A, 129 C, 136 A, 144 T, 189 C, 237 T, 246 C, 264 A, 327 C, 348 T, 357 C, 363 T, 387 A, 392 T, 502 C, 573 C [Hosts: Crambidae, Eulepte concordalis] Apanteles felipechavarriai Fernández-Triana, sp. n. (N = 1)
Barcoding region with 18 diagnostic nucleotides at positions: 81 A, 82 T, 99 T, 129 T, 136 T, 144 A, 189 T, 237 C, 246 T, 264 T or C, 327 T, 348 C, 357 T, 363 A, 387 T, 392 A or C, 502 T, 573 A or T 4
4(3) Ovipositor sheaths 1.4 × as long as metatibia (Fig. 23c); T1 length at most 1.9 × its width at posterior margin [Hosts: Tortricidae, Episimus sp.; Yponomeutidae, Atteva zebra] Apanteles adrianachavarriae Fernández-Triana, sp. n.
Ovipositor sheaths at most 1.2 × as long as metatibia; T1 length at least 2.1 × its width at posterior margin 5
5(4) Ovipositor sheaths length 0.8–0.9 × metatibia length (Fig. 30a); T2 width at posterior margin at most 3.7 × its length; body length 2.8 mm; fore wing length 2.8 mm [Hosts: Crambidae, Pilocrocis xanthozonalis, Tortricidae, Amorbia productana] Apanteles ronaldquirosi Fernández-Triana, sp. n. (N = 3)
Ovipositor sheaths length 1.0–1.2 × metatibia length (Figs 27c, 28a); T2 width at posterior margin at least 3.8 × its length; body length 2.2–2.4 mm (rarely 2.5 mm); fore wing length 2.4–2.6 mm 6
6(5) Fore wing with vein r 1.7 × as long as vein 2RS; flagellomerus 2 2.9 × as long as wide; flagellomerus 14 1.7 × as long as wide [Hosts: Crambidae, Asturodes fimbriauralis] Apanteles irenecarrilloae Fernández-Triana, sp. n. (N = 2)
Fore wing with vein r at most 1.4 × as long as vein 2RS; flagellomerus 2 3.1 × as long as wide; flagellomerus 14 at most 1.5 × as long as wide [Hosts: Crambidae, Diacme sp.] Apanteles luiscantillanoi Fernández-Triana, sp. n. (N = 3)
7(2) Ovipositor sheaths at most 0.8 × metatibia length (Figs 25a, d) [Hosts: Yponomeutidae, Atteva spp.] Apanteles anamartinesae Fernández-Triana, sp. n.
Ovipositor sheaths at least 1.0 × metatibia length (Figs 24a, b, 31a, c) 8
8(7) T1 length 1.7 × its width at posterior margin; T2 width at posterior margin 4.4 × its length [Hosts: Elachistidae, Antaeotricha similis, Stenoma sp.] Apanteles adrianguadamuzi Fernández-Triana, sp. n. (N = 2)
T1 length 1.5 × its width at posterior margin; T2 width at posterior margin 5.2 × its length [Hosts: Tortricidae, Episimus spp.] Apanteles yilbertalvaradoi Fernández-Triana, sp. n. (N = 2)
adrianaguilarae species-group

This group comprises three species characterized by extensive yellow-orange coloration, ocular-ocellar line 2.5 × posterior ocellus diameter, and fore wing with vein 2M as long as vein (RS+M)b. The group is strongly supported by the Bayesian molecular analysis (PP: 1.0, Fig. 1). Hosts: Tortricidae. All the described species are from ACG.

Key to species of the adrianaguilarae group
1 Ovipositor sheaths 0.9–1.0 × metatibia length (Figs 33a, c); fore wing with vein r 1.1 × as long as vein 2RS, vein 2RS 2.0 × as long as vein 2M, and vein 2M 0.7 × as long as vein (RS+M)b; pterostigma 3.6 × as long as wide; metafemur at least 3.1 × as long as wide Apanteles ivonnetranae Fernández-Triana, sp. n.
Ovipositor sheaths at most 0.6 × metatibia length (Figs 32d, 34c); fore wing with vein r at least 1.4 × as long as vein 2RS, vein 2RS at most 1.2 × as long as vein 2M, and vein 2M at least 1.0 × as long as vein (RS+M)b; pterostigma at most 3.1 × as long as wide; metafemur at most 2.9 × as long as wide 2
2(1) Metafemur mostly yellow, at most brown on posterior 0.3 (usually less) (Figs 32a, d); interocellar distance 2.2 × posterior ocellus diameter; T2 width at posterior margin 4.5 × its length; fore wing with vein 2RS 1.2 × vein 2M Apanteles adrianaguilarae Fernández-Triana, sp. n.
Metafemur mostly brown, at most yellow on anterior 0.4 (usually less) (Figs 34a, d); interocellar distance 1.8 × posterior ocellus diameter; T2 width at posterior margin 3.7 × its length; fore wing with vein 2RS 0.9 × vein 2M Apanteles vannesabrenesae Fernández-Triana, sp. n.
alejandromorai species-group

This group comprises 13 species which are unique among all Mesoamerican Apanteles in having an almost quadrate mediotergite 2 and a very long ovipositor. Both the Bayesian and neighbour joining trees (Figs 1, 2) have the species of this group in two separate clusters, each of them strongly supported (PP: 0.99 and 1.0 respectively, Fig. 1). Whenever the wasp biology is known, all are solitary parasitoids, with individual, white cocoons attached to the leaves where the caterpillar was feeding. Hosts: Elachistidae and Gelechiidae. All described species are from ACG, although we have seen undescribed species from other Neotropical areas.

Key to species of the alejandromorai group
1 Meso- and metafemora yellow (metafemora may have small, dark spot on posterior 0.1); metatibia mostly yellow, at most with dark brown to black spot in posterior 0.2 or less (rarely 0.3) of its length (Figs 39a, c, g, 42a, c, 45a) 2
Mesofemur (partially or completely) and metafemur (completely) dark brown to black; metatibia usually brown to black in posterior 0.3-0.5 (rarely 0.2) of its length (Figs 38a, c, e, 40a, c, 41a, c, 43a, c, 44a, 46a, 47a, c, 48a, 49a, c, 50a, c) 4
2(1) Ovipositor sheaths 1.2 × metatibia length (Figs 42a, c); body and fore wing length at most 3.2 mm; ocular-ocellar line 2.6 × posterior ocellus diameter; interocellar distance 2.2 × posterior ocellus diameter [Hosts: Elachistidae, Antaeotricha] Apanteles franciscoramirezi Fernández-Triana, sp. n. (N = 1)
Ovipositor sheaths at least 1.7 × metatibia length (Figs 39a, c, 45a, c); body and fore wing length at least 3.4 mm; ocular-ocellar line at most 1.9 × posterior ocellus diameter; interocellar distance at most 1.9 × posterior ocellus diameter; terostigma completely dark brown (at most with small pale spot at base); most of fore wing veins brown 3
3(2) Ovipositor sheaths 1.8 mm long; fore wing length 1.9 × as long as ovipositor sheaths length [Hosts: Antaeotricha radicalis and other Elachistidae feeding on Melastomataceae] Apanteles deifiliadavilae Fernández-Triana, sp. n. (N = 1)
Ovipositor sheaths 2.1–2.3 mm long; fore wing length 1.6–1.7 × as long as ovipositor sheaths length [Host: Antaeotricha spp. ] Apanteles juancarrilloi Fernández-Triana, sp. n. (N = 5)
4(1) All trochantelli, profemur, tegula and humeral complex entirely yellow (Figs 49a, c, g); mesofemur partially yellow, especially dorsally; metafemur white to yellow on anterior 0.1–0.2, giving the appareance of a light anellus (Fig. 49c) Apanteles tiboshartae Fernández-Triana, sp. n.
All trochantelli and part of profemur (basal 0.2–0.5) dark brown to black, tegula yellow, humeral complex half brown, half yellow; meso- and metafemur completely dark brown to black (mesofemur rarely with 0.2 or less yellow) 5
5(4) Ovipositor sheaths at most 1.6 × as long as metatibia length 6
Ovipositor sheaths at least 1.8 × as long as metatibia length 9
6(5) Pterostigma mostly dark brown with small, paler area centrally (Fig. 44b); T1 length at least 3.0 × its width at posterior margin Apanteles gabrielagutierrezae Fernández-Triana, sp. n. (N = 2)
Pterostigma mostly pale (yellow-white) or transparent, with only thin borders brown (Figs 43b, 46b, 47b); T1 length at most 2.8 × its width at posterior margin 7
7(6) Body length and fore wing length 3.0 mm; T1 width at posterior margin 0.6 × width at anterior margin [Hosts: Choreutidae, Tortyra; Elachistidae, Anacampsis] Apanteles luisgarciai Fernández-Triana, sp. n. (N = 1)
Body length and fore wing length at least 3.3 mm; T1 width at posterior margin 0.8 × width at anterior margin [Hosts: Elachistidae, Antaeotricha spp.] 8
8(7) Scutoscutellar sulcus with 8 pits; fore wing with vein r 2.2 × vein 2RS, and vein 2RS 1.3 × vein 2M; T1 length 2.7 × its width at posterior margin; flagellomerus 2 2.9 × as long as wide; flagellomerus 14 1.8 × as long as wide; ocular-ocellar line 2.3 × posterior ocellus diameter Apanteles luisbrizuelai Fernández-Triana, sp. n. (N = 1)
Scutoscutellar sulcus with at least 11 pits; fore wing with vein r 1.4 × vein 2RS, and vein 2RS 1.6 × vein 2M; T1 length 2.3 × its width at posterior margin; flagellomerus 2 2.6 × as long as wide; flagellomerus 14 1.5 × as long as wide; ocular-ocellar line 2.6 × posterior ocellus diameter Apanteles freddysalazari Fernández-Triana, sp. n. (N = 2)
9(5) Pterostigma mostly dark brown with small, paler area centrally (Fig. 38b); fore wing with vein 2RS 1.9 × vein 2M; flagellomerus 2 3.0 × as long as wide Apanteles alejandromorai Fernández-Triana, sp. n.
Pterostigma mostly pale (yellow-white) or transparent, with only thin borders brown (Figs 40b, 41b, 48b, 50b); fore wing with vein 2RS at most 1.6 × vein 2M (usually much less); flagellomerus 2 at most 2.8 × as long as wide 10
10(9) Metatibia mostly orange, with posterior 0.2 light brown (Figs 40a, c); flagellomerus 14 2.0 × as long as wide [Elachistidae] Apanteles eulogiosequeirai Fernández-Triana, sp. n. (N = 1)
Metatibia with posterior 0.4–0.5 dark brown to black (Figs 41c, 48a, 50c); flagellomerus 14 at most 1.7 × as long as wide [Elachistidae] 11
11(10) T1 length 2.2 × its width at posterior margin; T2 width at posterior margin 2.2 × its length; metafemur 3.2–3.3 × as long as wide [Elachistidae] Apanteles minornavarroi Fernández-Triana, sp. n.
T1 length at least 2.4 × its width at posterior margin; T2 width at posterior margin at most 1.9 × its length; metafemur 2.9–3.1 × as long as wide [Elachistidae] 12
12(11) T1 length 2.4 × its width at posterior margin; fore wing with vein r at least 2.3 × vein 2RS, vein 2RS 1.1 × vein 2M, and pterostigma 3.2 × as long as wide [Elachistidae] Apanteles marvinmendozai Fernández-Triana, sp. n. (N = 1)
T1 length 2.9 × its width at posterior margin; fore wing with vein r 1.8 × vein 2RS, vein 2RS 1.5 × vein 2M, and pterostigma 3.8 × as long as wide [Elachistidae] Apanteles fernandochavarriai Fernández-Triana, sp. n. (N = 4)
anabellecordobae species-group

This group comprises 14 species and is defined by the hypopygium either unfolded or with a relatively wide and translucid fold with none or very few (1-3) pleats only in the outermost area of fold. The species have a thick ovipositor (as thick as or thicker than width of median flagellomerus), with anterior width 3.0-5.0 × its posterior width beyond the constriction. The group is strongly supported by the Bayesian molecular analysis (PP: 1.0, Fig. 1). Hosts: Hesperiidae: Eudaminae, Hesperiinae, and Pyrginae; mostly gregarious parasitoids of leaf-rolling caterpillars (only two species are solitary parasitoids, with molecular data suggesting they form a sub-group on its own). All described species are from ACG, although we have seen numerous undescribed species from other Neotropical areas.

Key to species of the anabellecordobae group
1 Hypopygium without a median fold, with 0 or, at most, 1 small pleat visible (Figs 51c, 54c, 56c, 63c) 2
Hypopygium with a median fold and a few (1–3) pleats visible (Figs 52c, 55c, 57c, 58c, 59c, 64c) 6
2(1) Meso and metafemur (completely), and metatibia (at least partially) dark brown to black (Fig. 51a); fore wing with pterostigma mostly brown (Fig. 51b); ovipositor sheaths at least 0.8 × as long as metatibia length (Figs 51a, c); T2 width at posterior margin 3.1 × its length [Hosts: Hesperiidae, Achlyodes spp.; hosts feeding on Rutaceae] Apanteles anabellecordobae Fernández-Triana, sp. n.
All femora and tibiae yellow (at most with some infuscation on posterior 0.2 × or less of metafemur and metatibia) (Figs 54a, 56a, 60a, 63a); fore wing pterostigma either mostly pale or transparent with thin brown borders or brown with pale area centrally (Figs 54b, 56b, 60b, 63b); ovipositor sheaths at most 0.7 × as long as metatibia length (usually smaller) (Figs 54a, c, 56a, 63a, c); T2 width at posterior margin at least 3.3 × its length [Hosts: Hesperiidae, Astraptes spp., Gorgythion begga pyralina and Sostrata bifasciata nordica; hosts feeding on Fabaceae, Malpighiaceae, Malvaceae, and Sapindaceae] 3
3(2) Metafemur and metatibia yellow to light brown, with posterior 0.2 × dark brown; tegula pale, humeral complex half pale, half dark; pterostigma brown, with small pale area centrally (Figs 54b, 63b) [Hosts: Hesperiidae, Eudaminae; hosts feeding on Fabaceae, Malvaceae, and Sapindaceae] 4
Metafemur, metatibia, tegula and humeral complex yellow; pterostigma mostly pale or transparent with thin brown borders (Figs 56b, 60b) [Hosts: Hesperiidae, Pyrginae; hosts feeding on Malpighiaceae] 5
4(3) Flagellomerus 2 2.6 × as long as wide; flagellomerus 14 1.9 × as long as wide; mesoscutellar disc 1.5 × as long as wide; T1 3.4 × as long as wide at posterior margin [Hosts: Hesperiidae, Astraptes spp.; hosts feeding on Fabaceae, Malvaceae, and Sapindaceae] Apanteles osvaldoespinozai Fernández-Triana, sp. n.
Flagellomerus 2 2.9 × as long as wide; flagellomerus 14 1.6 × as long as wide; mesoscutellar disc 1.2 × as long as wide; T1 2.7 × as long as wide at posterior margin [Hosts: Hesperiidae, Astraptes spp.; hosts feeding on Fabaceae] Apanteles edwinapui Fernández-Triana, sp. n.
5(3) Pro- and mesocoxae dark brown, metacoxa black; flagellomerus 2 2.2 × as long as wide; T2 width at posterior margin 3.6 × its length [Host: Hesperiidae, Gorgythion begga pyralina feeding on Malpighiaceae deep into rainforests] Apanteles luciariosae Fernández-Triana, sp. n.
Pro- and mesocoxae yellow-brown, metacoxa dark brown; flagellomerus 2 3.0 × as long as wide; T2 width at posterior margin 4.7 × its length [Host: Hesperiidae, Gorgythion begga pyralina and Sostrata bifasciata nordica, feeding on Malpighiaceae in dry and rainforests] Apanteles freddyquesadai Fernández-Triana, sp. n.
6(1) T1 almost completely smooth and polished, at most with few punctures near posterior margin (Fig. 62g); propodeal areola with longitudinal carinae strongly converging posteriorly, running closely parallel (almost fused) for the posterior third of propodeum length until reaching nucha (Fig. 62g) [Hosts: Hesperiidae, Polythrix kanshul] Apanteles marianopereirai Fernández-Triana, sp. n.
T1 with at least some sculpture in posterior 0.3-0.5 (Figs 52e, 53f, 57f, 58f, 59f, 61f, 64h); propodeal carina with longitudinal carinae converging right before reaching nucha, not running closely parallel (Figs 52e, 53f, 57f, 58f, 59f, 61f, 64h) 7
7(6) Meso- and metafemora entirely or mostly dark brown to black (Figs 59a, c) [Host: Hesperiidae, Noctuana lactifera] Apanteles joseperezi Fernández-Triana, sp. n.
All femora mostly yellow (sometimes a small dark spot present on posterior end of metafemur), or mesofemur yellow and metafemur brown dorsally and yellow ventrally (Figs 52a, 53a, c, 55a, c, 57a, 58a, 61a, 64a) 8
8(7) Metasoma almost completely yellow (Figs 61a, c, f), except for T1 and T2 (males may have metasoma brown, if so then T3+ paler than T1-T2) [Hosts: Hesperiidae, Eudaminae, Telemiades antiope] Apanteles manuelpereirai Fernández-Triana, sp. n.
Metasoma mostly dark brown to black, the yellow parts, if any, limited to some sternites and/or laterotergites [Hosts: Hesperiidae, Pyrginae] 9
9(8) Pterostigma brown with at most a small pale spot at base, most veins brown (Figs 53b, 57b, 64b) 10
Pterostigma transparent or whitish with only thin brown borders, most veins transparent (Figs 52b, 55b, 58b) 12
10(9) T1 3.0 × as long as wide at posterior margin (Fig. 57f); antenna about same length than body; flagellomerus 14 1.4 × as long as wide; metatibial inner spur 1.5 × as long as metatibial outer spur; fore wing with vein r 2.0 × as long as vein 2RS [Host: Hesperiidae, Nisoniades godma] Apanteles guillermopereirai Fernández-Triana, sp. n.
T1 at least 3.6 × as long as wide at posterior margin (Fig. 64h); antenna clearly shorter than body; flagellomerus 14 at most 1.2 × as long as wide; metatibial inner spur at least 1.8 × as long as metatibial outer spur; fore wing with vein r 1.6 × as long as vein 2RS [Hosts: Hesperiidae, Staphylus spp.] 11
11(10) Metafemur, metatibia and metatarsus yellow, at most with small dark spots in apex of metafemur and metatibia (Fig. 64a) [Hosts: Hesperiidae, Staphylus vulgata] Apanteles ruthfrancoae Fernández-Triana, sp. n.
Metafemur brown dorsally and yellow ventrally, metatibia with a darker area on apical 0.2–0.3 ×, metatarsus dark (Figs 53a, c) [Hosts: Hesperiidae, Staphylus evemerus] Apanteles duniagarciae Fernández-Triana, sp. n.
12(9) T1 at least 4.0 × as long as posterior width (Fig. 55f); flagellomerus 14 2.3 × as long as wide; flagellomerus 2 1.6 × as long as flagellomerus 14; metafemur 3.3 × as long as wide; mesocutum and mesoscutellar disc mostly heavily and densely punctured; body length 3.3–3.6 mm and fore wing length 3.3–3.6 mm [Hosts: Hesperiidae, Pyrrhopyge zenodorus] Apanteles eldarayae Fernández-Triana, sp. n.
T1 at most 2.6 × as long as posterior width (Figs 52e, 58f); flagellomerus 14 at most 1.4 × as long as wide; flagellomerus 2 at least 2.0 × as long as flagellomerus 14; metafemur at most 3.0 × as long as wide; mesocutum and mesoscutellar disc mostly smooth or with sparse, shallow punctures; body length 2.4–2.6 mm and fore wing length 2.5–2.7 mm 13
13(12) T2 width at posterior margin 3.6 × its length; fore wing with vein r 2.4 × as long as vein 2RS, and vein 2RS 0.9 × as long as vein 2M [Hosts: Hesperiidae, Timochreon satyrus, Anisochoria polysticta] Apanteles harryramirezi Fernández-Triana, sp. n.
T2 width at posterior margin 4.3 × its length; fore wing with vein r 1.6 × as long as vein 2RS, and vein 2RS 1.5 × as long as vein 2M [Hosts: Hesperiidae, Pyrgus spp., Heliopetes arsalte] Apanteles carolinacanoae Fernández-Triana, sp. n.
anamarencoae species-group

This group comprises two species, characterized by pterostigma fully brown; all coxae dark brown to black; tegula, humeral complex, all femora and all tibiae yellow (metafemur with small brown spot on posterior 0.2 × or less); and ovipositor sheaths at least 1.4 × as long as metatibia length. Molecular data does not support this group. Hosts: Tortricidae, Elachistidae, Oecophoridae. All described species are from ACG.

Key to species of the anamarencoae species-group
1 Scape anterior 0.6–0.7, entire metatibia and metatarsus yellow (Figs 66a, c, e) [Hosts: Tortricidae] Apanteles juanlopezi Fernández-Triana, sp. n. (N = 2)
Scape almost completely dark brown (Fig. 65d); metatibia with small dark spot on posterior 0.1 ×; metatarsus with segment 1 brown to dark brown on posterior 0.5–0.6, remaining segments with some brown marks (Figs 65a, c) [Hosts: Elachistidae, Oecophoridae] Apanteles anamarencoae Fernández-Triana, sp. n. (N = 3)
arielopezi species-group

This group comprises two species, characterized by relatively small body size (body length at most 2.4 mm and fore wing length at most 2.7 mm), mesoscutellar disc smooth, tegula and humeral complex of different color, and brown pterostigma. The group is strongly supported by the Bayesian molecular analysis (PP: 1.0, Fig. 1). Hosts: Tortricidae, Elachistidae. All described species are from ACG.

Key to species of the arielopezi group
1 Antenna shorter than body length, extending to half metasoma length; ovipositor sheaths slightly shorter (0.9 ×) than metatibia length (Figs 69a, c) Apanteles arielopezi Fernández-Triana, sp. n.
Antenna about same length than body; ovipositor sheaths 1.3 × as long as metatibia length (Figs 70a, c) Apanteles mauriciogurdiani Fernández-Triana, sp. n.
ater species-group

Proposed by Nixon, this is a heterogeneous assemble that contains “many aggregates of species that are not closely related but merge into one another through transitional forms”, and is characterized by having “a well defined areola and costulae in the propodeum, and a vannal lobe that is centrally concave and without setae” (Nixon 1965: 25). Such a general and vague definition created a largely artificial group, including many species worldwide (e.g., Nixon 1965; Mason 1981). Known hosts for the ater species-group vary considerably, and the molecular data available for some species (Figs 1, 2) does not support this group either. Future study of the world fauna will likely split the group into smaller, better defined units. For the time being, and just for Mesoamerica, we are keeping here three previously described species (Apanteles galleriae, Apanteles impiger and Apanteles leucopus), as well as six new species that do not fit into any of the other species-groups considered for the region which keeps this as a “garbage can” group. Another six previously described Apanteles with Mesoamerican distribution which used to be part of the ater group are here removed from that group and transferred as follows: Apanteles carpatus to the newly created carpatus species-group, Apanteles leucostigmus to the newly created leucostigmus group, Apanteles megathymi to the newly created megathymi species-group, Apanteles paranthrenidis and Apanteles thurberiae to the newly created paranthrenidis group, and Apanteles vulgaris to the newly created vulgaris species-group.

Key to species of the ater species-group

[The species Apanteles leucopus is placed in the ater species-group but we could not study any specimens, just photos of the holotype sent from the BMNH (Fig. 78). Unfortunately, the illustrations do not provide all details needed to include the species in any key of this paper]

1 Pterostigma relatively broad, its length less than 2.5 × its width Apanteles galleriae Wilkinson, 1932
Pterostigma relatively narrow, its length more than 3.0 × its width 2
2(1) Pterostigma entirely brown or brown with pale spot at base (Figs 72b, 73b, 74b, 76b, 77b) 2
Pterostigma entirely transparent or mostly transparent with only thin brown borders (as in Fig. 71b) 7
3(2) Tarsal claws simple Apanteles josejaramilloi Fernández-Triana, sp. n. (N = 1)
Tarsal claws with a single basal spine-like seta 4
4(3) Metacoxa entirely dark brown to black (Fig. 74b); scutoscutellar sulcus thin and with more than 10 close and small impressed pits Apanteles franciscopizarroi Fernández-Triana, sp. n. (N = 1)
Metacoxa entirely yellow-white or orange, at most with small brown spot on anterior end (Figs 72a, c, 73a, c, f, 76a); scutoscutellar sulcus relatively wide, with at most 7 widely impressed pits 5
5(4) Mesoscutellar disc mostly smooth; T2 and T3 yellow-orange (Fig. 76f) Apanteles jairomoyai Fernández-Triana, sp. n. (N = 1)
Mesoscutellar disc mostly punctured; T2 and T3 black (Figs 72g, 73f) 6
6(5) Mesocoxa yellow with anterior 0.3 brown (Fig. 72a); antenna dark brown to black (Figs 72d-f); labrum and tegula dark brown (Figs 72f, g); stigma brown; body length 2.3 mm, and fore wing length 2.6 mm; T1 3.5 × as long as wide; T2 with some sculpture on posterior margin Apanteles cristianalemani Fernández-Triana, sp. n. (N = 1)
Mesocoxa entirely yellow (Fig. 73a); antenna with scape and pedicel yellow (Figs 73d, e); labrum yellow (Fig. 73e), tegula yellow-white (Fig. 73f); stigma brown with small pale spot at base; body length 3.7 mm, and fore wing length 3.7 mm; T1 2.4 × as long as wide; T2 smooth Apanteles diegoalpizari Fernández-Triana, sp. n. (N = 4)
7(2) Pro-, meso-, and part of metacoxa yellow-orange; tegula and humeral complex yellow (Fig. 75g) Apanteles impiger Muesebeck, 1958
At least meso- and metacoxae (sometimes also procoxa) dark brown to black (Figs 71a, g); tegula and humeral complex dark brown to black (Fig. 71g) Apanteles anariasae Fernández-Triana, sp. n. (N = 1)
bernyapui species-group

This group comprises four species, characterized by extensive yellow coloration (and usually orange marks on posterior 0.2–0.3 × of anteromesoscutum and upper anterior corner of mesopleura), T1 black (same color of propodeum) and mostly strongly sculptured, with longitudinal striation laterally and a central excavated area with transverse striation. The group is strongly supported by the Bayesian molecular analysis (PP: 1.0, Fig. 1). Hosts: mostly Crambidae, with some records from Elachistidae, Gelechiidae and Noctuidae. All described species are from ACG.

Key to species of the bernyapui group
1 Anteromesoscutum and mesopleura completely black (Figs 79a, g) Apanteles bernyapui Fernández-Triana, sp. n.
Anteromesoscutum with posterior 0.2–0.3 (especially centrally and along posterior margin) and upper anterior corner of mesopleura orange (Figs 80f, 82g) 2
2(1) Body length 2.3–2.4 mm; fore wing length 2.5–2.6 mm; ovipositor sheaths 0.6 × as long as metatibia; fore wing with vein r 1.7 × as long as vein 2RS; mesoscutellar disc rather strongly punctured near margins (Fig. 82g) Apanteles victorbarrantesi Fernández-Triana, sp. n. (N = 4)
Body length length at least 2.7 mm (usually more); fore wing length at least 2.9 mm (usually more); ovipositor sheaths at least 0.8 × as long as metatibia; fore wing with vein r at most 1.4 × as long as vein 2RS; mesoscutellar disc either smooth, or with shallow punctures (Figs 80f, 81g) 3
3(2) T1 2.3 × as long as wide at posterior margin; T2 3.9 × as wide as its medial length (Fig. 81g); ovipositor sheaths shorter (0.8 ×) than metatibia; mesoscutellar disc mostly smooth; mesofemur mostly light yellow, with posterior 0.1 light orange; metatibia with anterior 0.6 light yellow, posterior 0.4 orange; ocular-ocellar line 2.0 × as long as posterior ocellus diameter; interocellar distance 1.7 × as long as posterior ocellus diameter; second flagellomerus 2.4 × as long as wide; metafemur 2.9 × as long as wide Apanteles raulacevedoi Fernández-Triana, sp. n.
T1 3.3 × as long as wide at posterior margin; T2 3.3 × as wide as its median length (Fig. 80f); ovipositor sheaths same length (1.0 ×) as metatibia; mesoscutellar disc with shallow punctures; mesofemur mostly yellow, with posterior 0.1–0.2 × dark brown; metatibia yellow, with posterior 0.3 dark brown; ocular-ocellar line 2.7 × as long as posterior ocellus diameter; interocellar distance 2.2 × as long as posterior ocellus diameter; second flagellomerus 3.0 × as long as wide; metafemur 3.3 × as long as wide Apanteles javiersihezari Fernández-Triana, sp. n. (N = 3)
bienvenidachavarriae species-group

This group comprises three species, sharing with the adelinamoralesae species-group similar morphological and biological (hosts) traits. They differ from the latter group in having meditergite 2 much less transverse, its width at posterior margin usually 2.5 × (at most 2.7 ×) its length – mediotergite 2 usually much more than 2.9 × in the adelinamoralesae species-group. The group is strongly supported by the Bayesian molecular analysis (PP: 1.0, Fig. 1); the single exception being Apanteles marisolarroyoae, which is included here interimly – its barcode does not cluster with the other two species although it shares with them morphological and host traits. Hosts: Elachistidae. All described species are from ACG.

Key to species of the bienvenidachavarriae group
1 Profemur except for at most anterior 0.2, mesofemur in posterior 0.2, and metatibia in anterior 0.7 orange-yellow (Figs 84a, c); antenna as long as body; larger species, body length 3.8–4.0 mm and fore wing length 3.9–4.0 mm [Hosts: Elachistidae, Anadasmus spp.] Apanteles bienvenidachavarriae Fernández-Triana, sp. n.
Promefur in anterior 0.5, mesofemur entirely, and metatibia in posterior 0.4–0.8 black to dark brown (Figs 85a, e, 86a, c); antenna shorter than body; smaller species, body length 3.0–3.3 mm and fore wing length 3.1–3.3 mm 2
2(1) Metatibia almost completely black, except for anterior 0.2 or less which is yellow; T1 2.6 × as long as wide at posterior margin [Hosts: Elachistidae, undetermined species] Apanteles marisolarroyoae Fernández-Triana, sp. n.
Metatibia at most with black on posterior 0.4–0.5; T1 2.3 × as long as wide at posterior margin [Hosts: Elachistidae, Antaeotricha zelleri, Gonioterma anna] Apanteles josecalvoi Fernández-Triana, sp. n. (N = 2)
calixtomoragai species-group

This group comprises three species with pectinate tarsal claws, an almost unique feature within the Mesoamerican species of Apanteles (the only two other species in the region known to have pectinate tarsal claws, Apanteles juliodiazi and Apanteles waldymedinai, can be easily separated based on its orange heads). Also, the calixtomoragai group contains the largest Apanteles in the region (+4.0 mm of body length). The group is strongly supported by the Bayesian molecular analysis (PP: 1.0, Fig. 1). All species are solitary, with the individual coccon (mostly white, but with basal 0.3–0.4 light brown) attached to the leaves where the caterpillar rests when not feeding. Hosts: Hesperiidae. All described species are from ACG, although we have seen undescribed species from other Neotropical areas.

Key to species of the calixtomoragai group
1 Sternites and hypopygium dark brown to black (Fig. 89a); all femora dark orange to reddish (Figs 89a, d); fore wing with apical 0.3–0.4 (beyond veins r and 2RS) slightly infumated, clearly darker than rest of wing (Fig. 89b); T1 and T2 with some sculpture near lateral and/or posterior margins (Fig. 89h); fore wing with vein 2RS 1.4 × as long as vein 2M; flagellomerus 14 2.7 × as long as wide (rarely up to 2.8 ×); body length usually over 4.7 mm (range: 4.4–5.2 mm); fore wing length 5.2–5.4 mm; mesoscutellum lunules 0.6–0.7 × as high as maximum height of lateral face of mesoscutellum [Hosts: Ouleus dilla baru] Apanteles petronariosae Fernández-Triana, sp. n.
Sternites and hypopygium mostly to completely yellow, at most light brown (as in Fig. 88a); pro- and mesofemora yellow, metafemur yellow or orange to reddish; fore wing mostly hyaline (if there is some infumation, it is very slightly and not restricted to wing apex) (Figs 87b, 88b); T1 and T2 mostly smooth (as in Fig. 87e); fore wing with vein 2RS 1.7–1.8 × as long as vein 2M; flagellomerus 14 2.8–3.1 × as long as wide; body length usually less than 4.5 mm (range: 4.0–4.9 mm); forewing length 4.5–5.1 mm; mesoscutellum lunules 0.4–0.5 × as high as maximum height of lateral face of mesoscutellum [Hosts: Milanion marciana and Quadrus cerialis] 2
2(1) Mesoscutellum with non-polished area of lateral face with striae interrupted dorsally by a smooth area marking a clear separation from axilla (axilla also with striated sculpture) (Fig. 87e); fore wing length usually 4.8 mm or less (range: 4.5–4.9 mm); body length 4.3 mm (range: 4.0–4.7 mm) [Hosts: Milanion marciana. A total of 22 diagnostic characters in the barcoding region: 67 T, 124 T, 133 C, 139 C, 181 T, 194 T, 200 C, 278 C, 298 T, 300 G, 311 A, 319 T, 335 G, 340 C, 346 C, 347 C, 523 T, 595 C, 616 C, 628 T, 634 C, 640 T] Apanteles calixtomoragai Fernández-Triana, sp. n.
Mesoscutellum with non-polished area of lateral face with striae that continue towards axilla, with no clear or polished area separating both striated surfaces (Fig. 88g); fore wing length almost always 5.0 mm or more (range: 4.8–5.1 mm); body length 4.5 mm (range: 4.1–4.9 mm) [Hosts: Quadrus cerialis. A total of 22 diagnostic characters in the barcoding region: 67 C, 124 C, 133 T, 139 T, 181 A, 194 C, 200 T, 278 T, 298 A, 300 A, 311 G, 319 A, 335 A, 340 T, 346 T, 347 T, 523 C, 595 T, 616 T, 628 A, 634 T, 640 C] Apanteles manuelriosi Fernández-Triana, sp. n.
carlosguadamuzi species-group

This group comprises six species with extensive yellow-orange coloration, smooth mesoscutellar disc, mediotergite 1 weakly sculptured and light coloured with orange-yellow to light brown (males tend to have tergites with darker coloration, compared to females). The group is strongly supported by the Bayesian molecular analysis (PP: 1.0, Fig. 1). Hosts: mostly Crambidae, but some species reared from Choreutidae, Elachistidae, and Gelechiidae. Some species are gregarious and some are solitary parasitoids. All described species are from ACG, although we have seen undescribed species from other Neotropical areas.

Key to species of the carlosguadamuzi group
1 T1 light brown, distinctly darker than T2 (Figs 91g, 93f) [Host: Ategumia lotanalis] 2
T1 entirely orange or orange-yellow, same color as T2 (Figs 90g, 92f, 94f) 3
2(1) Fore wing with vein r 1.8–2.0 × as long as vein 2RS, and vein 2RS 1.0 × as long as vein 2M Apanteles cinthiabarrantesae Fernández-Triana, sp. n.
Fore wing with vein r 1.3 × as long as vein 2RS, and vein 2RS 1.6 × as long as vein 2M Apanteles javiercontrerasi Fernández-Triana, sp. n.
3(1) T2 width at posterior margin at most 3.1 × its median length (Fig. 94f); ocular-ocellar line at most 1.8 × posterior ocellus diameter 4
T2 width at posterior margin at least 3.9 × its median length (Figs 90g, 92f); ocular-ocellar line at least 2.1 × posterior ocellus diameter 5
4(3) T1 2.5 × as long as wide at posterior margin; T2 width at posterior margin 3.1 × median length; fore wing with vein 2RS 1.6 × as long as vein 2M [Hosts: Gelechiidae] Apanteles jesusbrenesi Fernández-Triana, sp. n. (N = 4)
T1 3.1 × as long as wide at posterior margin; T2 width at posterior margin 2.7 × median length; fore wing with vein 2RS 1.9 × as long as vein 2M [Hosts: Elachistidae] Apanteles williamcamposi Fernández-Triana, sp. n. (N = 2)
5(3) Metatarsus, posterior 0.3 of metatibia, and posterior 0.1 of metafemur brown to black, contrasting with rest of hind leg which is orange-yellow; body length 3.2–3.4 mm; fore wing length 3.4–3.6 mm; fore wing with vein r 2.1 × as long as 2RS; flagellomerus 2 2.6 × as long as wide; metafemur 3.2 × as long as wide [Hosts: Choreutidae, Crambidae] Apanteles carlosguadamuzi Fernández-Triana, sp. n. (N = 5)
Metatarsus yellow or orange-yellow, same color as rest of hind leg, except for 0.2 or less of metatibia which is brown; body length usually 2.5–2.7 mm (rarely up to 3.0 mm); fore wing length 2.7–2.9 mm (rarely up to 3.2 mm); fore wing with vein r 1.3 × as long as 2RS; flagellomerus 2 3.2 × as long as wide; metafemur 2.9 × as long as wide [Hosts: Crambidae] Apanteles edithlopezae Fernández-Triana, sp. n.
carlosrodriguezi species-group

This group comprises three species, characterized by hypopygium with relatively short fold where no pleats (or at most one weak pleat) are visible, ovipositor sheaths very short (0.4–0.5 × as long as metatibia), and relatively small size (body length and fore wing length not surpassing 2.5 mm). Another Mesoamerican species, Apanteles aidalopezae shares that combination of characters, but can be separate from the carlosrodriguezi species-group because of its white pterostigma, transparent or white fore wing veins, and rather elongate glossa. The group is strongly supported by the Bayesian molecular analysis for two of its three component species (PP: 0.99, Fig. 1), however, Apanteles carlosrodriguezi clusters apart and future studies may find it is better to split it. Morphological data (especially shape of hypopygium and ovipositor sheaths length) suggest that the species might be placed on a new genus on their own when the phylogeny of Microgastrinae is better resolved. Because that is beyond the scope of this paper, we describe the species under Apanteles – the best arrangement at the moment. Hosts: Mostly gregarious on Crambidae; but Apanteles carlosrodriguezi is a solitary parasitoid on Elachistidae and possible Choreutidae. All described species are from ACG.

Key to species of the carlosrodriguezi group
1 All coxae, most of metatibia, meso- and metafemora dark brown to black (Figs 96a, c, g); body length and fore wing length 1.9–2.0 mm [Solitary parasitoid] Apanteles carlosrodriguezi Fernández-Triana, sp. n. (N = 3)
All coxae except for posterior 0.5 of metacoxa, at least anterior 0.3 × of metatibia, most of meso- and metafemora, yellow or white-yellow (Figs 97a, c, 98a, c); body length and fore wing length at least 2.2 mm [Gregarious parasitoids] 2
2(1) Face reddish-brown, clearly different in color from rest of head, which is dark brown to black (Fig. 98d); metafemur entirely yellow or at most with brown spot dorsally on posterior 0.2–0.3 (Fig. 98c); metatibia brown on posterior 0.6–0.7 (Fig. 98a) [A total of 32 diagnostic characters in the barcoding region: 23 T, 37 G, 68 T, 74 C, 88 A, 181 T, 203 T, 247 C, 259 C, 271 T, 278 T, 295 C, 311 T, 328 A, 346 A, 359 C, 364 T, 385 T, 428 C, 445 C, 448 C, 451 T, 467 C, 490 C, 500 C, 531 C, 544 T, 547 T, 574 C, 577 T, 601 T, 628 A] Apanteles robertoespinozai Fernández-Triana, sp. n.
Face almost always dark brown to black, same color as rest of head (Fig. 97e); metafemur brown dorsally on posterior 0.5–0.8 (Fig. 97c); metatibia brown on posterior 0.4–0.5 (Fig. 97a, c) [A total of 32 diagnostic characters in the barcoding region: 23 C, 37 A, 68 C, 74 T, 88 G, 181 A, 203 C, 247 T, 259 T, 271 C, 278 C, 295 T, 311 G, 328 T, 346 T, 359 T, 364 A, 385 C, 428 T, 445 T, 448 T, 451 C, 467 T, 490 T, 500 T, 531 T, 544 A, 547 A, 574 T, 577 C, 601 C, 628 T] Apanteles gloriasihezarae Fernández-Triana, sp. n.
carloszunigai species-group

This group comprises two species, characterized by the combination of folded hypopygium with very few (usually 1-3) pleats occupying just outermost area of fold, small size (fore wing less than 2.8 mm), and all coxae completely yellow. The group is supported by the Bayesian molecular analysis (PP: 0.66, Fig. 1). No host is known for this species-group. All the described species are from ACG; we have seen another species from ACG which cannot be described here because of poor condition of its known specimen.

Key to species of the carloszunigai group
1 Metafemur and metatibia almost entirely orange, with light brown spot on posterior 0.1 × (Figs 100a, c); T1 length 3.7 × its width at posterior margin; T2 width at posterior margin 3.5 × its length (Fig. 100g); flagellomerus 2 2.6 × as long as wide Apanteles yeissonchavesi Fernández-Triana, sp. n. (N = 1)
Metafemur and metatibia with posterior 0.2–0.3 × brown (Figs 99a, c); T1 length 3.2 × its width at posterior margin; T2 width at posterior margin 4.0 × its length (Fig. 99g); flagellomerus 2 3.0 × as long as wide Apanteles carloszunigai Fernández-Triana, sp. n. (N = 1)
carpatus species-group

Until now, Apanteles carpatus had been placed within the ater species group. However, we found that the combination of a relatively broad pterostigma (its length less than 3.0 × its width) and mediotergite 2 mostly sculptured with strong longitudinal striation, seems to be characteristic of several Mesoamerican species; which are also strongly supported as a group by the Bayesian molecular analysis (PP: 0.99, Fig. 1). Thus, we here consider them as a distinct group, which so far comprises five species but it is likely to include more when other Neotropical areas are studied. The only hosts known are for Apanteles carpatus, a cosmopolitan species with nine different families of host recorded, many of them dubious. More study will be required before accurate host families associated with this species-group can be established.

Key to species of the carpatus group
1 T2 length at least 2.5 × its width at posterior margin (Figs 101f, 102g) 2
T2 length at most 1.6 × its width at posterior margin (Figs 103g, 104g) 3
2(1) Metacoxa with posterior 0.3 yellow (Fig. 102a); body length and fore wing length at most 2.2 mm; mesoscutellar disc mostly smooth (Fig. 102f); scutoscutellar sulcus with 11–12 impressions (Fig. 102f); ocular-ocellar line at most 1.8 × posterior ocellus diameter Apanteles rhomboidalis (Ashmead, 1900)
Metacoxa brown; body length and fore wing length at least 2.8 mm; mesoscutellar disc mostly sculptured or sculptured near margins (Fig. 101f); scutoscutellar sulcus with 5–6 impressions (Fig. 101f); ocular-ocellar line 2.4 × posterior ocellus diameter Apanteles albanjimenezi Fernández-Triana, sp. n. (N = 1)
3(2) Fore wing with vein 2RS 1.4 × vein 2M, and vein 2M 0.7 × vein (RS+M)b; body length usually 2.5–2.6 mm (rarely up to 2.8 mm) and fore wing length 2.6–2.7 mm (rarely up to 2.9 mm); anteromesoscutum with extensive orange coloration (Fig. 104g); metatibia inner spur 0.5 × metabasitarsus length Apanteles rolandoramosi Fernández-Triana, sp. n. (N = 4)
Fore wing with vein 2RS at most 1.1 × vein 2M, and vein 2M at least 0.9 × vein (RS+M)b; body length and fore wing length usually 3.0 mm or more (rarely less); anteromesoscutum black (Fig. 103g); metatibia inner spur at least 0.6 × metabasitarsus length 4
4(3) Flagellomerus 2 2.6 × as long as wide; flagellomerus 2 length 2.6 × flagellomerus 14 length; tarsal claws simple; T1 parallel-sided; metacoxa partially yellow (Fig. 103a); ocular-ocellar line 1.8 × posterior ocellus diameter Apanteles robertovargasi Fernández-Triana, sp. n. (N = 1)
Flagellomerus 2 at most 2.2 × as long as wide; flagellomerus 2 length at most 2.2 × flagellomerus 14 length; tarsal claws with single basal spine-like seta; T1 clearly widening towards posterior margin; metacoxa entirely brown; ocular-ocellar line at least 2.0 × posterior ocellus diameter Apanteles carpatus (Say, 1836)
coffeellae species-group

This is an artificial group, neither supported by molecular nor host data, but only for some morphological resemblance of the species. It comprises Apanteles coffeellae (the only described species of Apanteles in Mesoamerica known to parasitize leaf-mining Lepidoptera), as well as three new species from ACG described below. It is characterized by its small size (body length 1.6–2.2 mm, fore wing length 2.0–2.2 mm), and mediotergite 1 strongly narrowing posteriorly. The known hosts (only for Apanteles coffeellae) include members of the Lepidoptera families Gracillariidae and Lyonetiidae, but no hosts are known for the other species. The described species are from the Caribbean and ACG, although it is likely that there are more undescribed species from other Neotropical areas. Future study might find this group to contain species of Apanteles parasitoids of leaf-mining Lepidoptera.

Key to species of the coffeellae group
1 T1 smooth and more than 4.0 × as long as its posterior width (Fig. 106g); fore wing length at most 1.8 mm Apanteles coffeellae Muesebeck, 1958
T1 mostly sculptured and less than 4.0 × as long as its posterior width (Fig. 107f, 108f, 109f); fore wing length at least 2.0 mm 2
2(1) Ovipositor sheaths 1.2 × as long as metatibia (Fig. 108a, c); propodeal areola without transverse carinae extending to spiracle Apanteles lisabearssae Fernández-Triana, sp. n.
Ovipositor sheaths at most 0.6 × as long as metatibia (Figs 107a, c, 109a, c); propodeal areola with transverse carinae extending to spiracle (as in Fig. 107f) 3
3(2) Mesoscutellar disc mostly punctured (Fig. 107f); mesofemur yellow (Fig. 107c); metatibia mostly dark brown, except for anterior 0.3, which is yellow; ovipositor sheaths 0.6 × as long as metatibia Apanteles laurahuberae Fernández-Triana, sp. n.
Mesoscutellar disc smooth (Fig. 109f); mesofemur dark brown on anterior 0.5 × (Fig. 109c); metatibia mostly yellow, except for posterior 0.3, which is dark brown; ovipositor sheaths 0.4 × as long as metatibia Apanteles mariaguevarae Fernández-Triana, sp. n. (N = 2)
diatraeae species-group

This group was proposed by Austin and Dangerfield (1989). Those authors considered it a monophyletic group, with striking body modifications associated with specialized parasitism of stem-borers in confined places. They included ten species in the group (seven from the New World, two from Africa and one from the Oriental region). It differs from the morphologically similar guadaluperodriguezae group in the propodeum areola shape and the parasitization of stem-borer Crambidae. In Mesoamerica three species are included: Apanteles deplanatus, Apanteles diatraeae, and Apanteles fredi. They are characterized by small size (1.8–2.2 mm in length), body distinctly flattened dorsoventrally, propodeal areola elongate and parallel-sided, and very short antenna (length not surpassing posterior margin of tergite 1). They are all greagarious. Hosts: Crambidae. Distribution: pantropical (Austin and Dangerfield 1989).

Key to species of the diatraeae group
1 T1 less than 2.0 × as long as width at its posterior margin and sculptured on its anterior half; T2 width at posterior margin 3.0 × its length; fore wing with vein r 1.0 × as long as vein 2RS; metatibia with basal 0.3 yellow, rest brown; mesoscutellum lunules at least 0.8 × as high as maximum height of lateral face of scutellum [Host: Crambidae (Diatraea sp.). Distribution: Guatemala] Apanteles fredi Austin & Dangerfield, 1989
T1 more than 2 × as long as width at its posterior margin and either mostly sculptured or with some sculpture near the lateral borders and/or the apical 0.3–0.5 (Figs 203g, 204g); T2 width at posterior margin less than 2.0 × its length; fore wing with vein r 1.4 × as long as vein 2RS; metatibia dark brown; mesoscutellum lunules 0.6–0.7 × as high as maximum height of lateral face of scutellum (Figs 203g, 204g) 2
2(1) T1 usually less than 3 × as long as width at its posterior margin and heavily sculptured medially, with two strong, median longitudinal carinae on apical half (Fig. 204g); anteromesoscutum mostly smooth or with shallow sparse punctures, except for anterior 0.3 where punctures are deeper and/or denser (Fig. 204g); fore wing with vein 2RS 1.3 × as long as vein 2M; ovipositor sheaths 0.8 × metatibia length (Fig. 204a, c) [Hosts: Crambidae (Diatraea spp., Galleria mellonella). Distribution: several Caribbean islands, Central and South America, introduced into the US, France and India] Apanteles diatraeae Muesebeck, 1921
T1 usually more than 3 × as long as width at its posterior margin and mostly smooth (Fig. 203g); anteromesoscutum mostly smooth (Fig. 203g); fore wing with vein 2RS 0.8 × as long as vein 2M; ovipositor sheaths 0.6 × metatibia length (Fig. 203a, c) [Host: Crambidae (Diatraea spp.). Distribution: Mexico] Apanteles deplanatus Muesebeck, 1957
dickyui species-group

This group comprises two species, characterized by pterostigma mostly transparent with only thin brown borders, tegula and humeral complex yellow, all coxae dark brown to black, mediotergite 1 at least 2.4 × as long as wide at posterior margin, and mediotergite 2 mostly scultured. The group is strongly supported by the Bayesian molecular analysis (PP: 1.0, Fig. 1). Hosts: Unknown. The described species are from ACG.

Key to species of the dickyui group
1 Interocellar distance 2.1 × as long as ocellus diameter; ocular-ocellar line 2.4 × as long as posterior ocellus diameter; flagellomerus 2 2.8 as long as wide; fore wing with vein R1 4.0 × as long as distance between ends of veins R1 and 3RS Apanteles eduardoramirezi Fernández-Triana, sp. n.
Interocellar distance 1.3 × as long as ocellus diameter; ocular-ocellar line 2.0 × as long as posterior ocellus diameter; flagellomerus 2 3.3 as long as wide; fore wing with vein R1 6.0 × as long as distance between ends of veins R1 and 3RS Apanteles dickyui Fernández-Triana, sp. n. (N = 1)
erickduartei species-group

This group comprises five species, characterized by extensive extensive yellow-orange coloration (including tegula and humeral complex, parts of the axillar complex, most of laterotergites 1–4, all sternites, and hypopygium), mesoscutellar disc mostly punctured, and mediotergite 1 more than 2.3 × as long as wide. The group is strongly supported by the Bayesian molecular analysis (PP: 1.0, Fig. 1). The species are solitary parasitoids. Hosts: Crambidae. All the described species are from ACG.

Key to species of the erickduartei group
1 Ovipositor sheaths 0.5 × metatibia length (Figs 115a, c); fore wing with vein r 2.4 × vein 2RS; T1 length 2.3 × its width at posterior margin Apanteles luishernandezi Fernández-Triana, sp. n. (N = 4)
Ovipositor sheaths at least 0.8 × metatibia length (usually more) (Figs 113a, c, 114a, c, 116a, c, 117a, c); fore wing with vein r at most 1.7 × vein 2RS; T1 length at least 2.5 × its width at posterior margin (usually more) 2
2(1) T3 mostly yellow (except for thin brown border on anterior margin) (Fig. 117g); metafemur with anterior 0.3–0.4 yellow, rest brown (Figs 117a, c); flagellomerus 2 2.2 × as long as wide Apanteles ronaldcastroi Fernández-Triana, sp. n. (N = 2)
T3 either entirely dark brown or with extensive, dark brown, central band, covering 0.4–0.5 of tergite and running from anterior to posterior margins (Figs 113g, 114f, 116f); metafemur either almost entirely dark brown, at most with small yellow spot on anterior 0.1 (usually), or entirely yellow (rarely) (Figs 113a, c, 114a, c, 115a); flagellomerus 2 at least 2.5 × as long as wide 3
3(2) Ovipositor sheaths 0.8 × metatibia length (rarely up to 0.9 ×) (Fig. 116a, c); T1 strongly narrowing towards posterior margin (maximum width of tergite 1.7 × width at posterior margin) (Fig. 116f); T3 entirely dark brown (Fig. 116f); flagellomerus 2 2.5 × as long as wide; flagellomerus 2 length 2.2 × flagellomerus 14 length; ocular-ocellar line 2.3 × posterior ocellus diameter; interocellar distance 2.2 × posterior ocellus diameter Apanteles milenagutierrezae Fernández-Triana, sp. n.
Ovipositor sheaths 1.0-1.2 × metatibia length (as in Figs 114a, c); T1 not so strongly narrowing towards posterior margin (maximum width of tergite 1.2–1.5 × width at posterior margin) (Figs 113g, 114f); T3 partially yellow (Figs 113g, 114f); flagellomerus 2 at least 2.7 × as long as wide; flagellomerus 2 length at least 2.5 × flagellomerus 14 length; ocular-ocellar line at most 2.1 × posterior ocellus diameter; interocellar distance 1.9 × posterior ocellus diameter 4
4(3) T1 lenght 3.2 × its width at posterior margin; ocular-ocellar line 2.1 × posterior ocellus diameter; flagellomerus 2 2.9 × as long as wide Apanteles felixcarmonai Fernández-Triana, sp. n. (N = 3)
T1 length 2.5–2.8 × its width at posterior margin; ocular-ocellar line 1.8 × posterior ocellus diameter; flagellomerus 2 2.7 × as long as wide Apanteles erickduartei Fernández-Triana, sp. n.
glenriverai species-group

This group contains two species characterized by its pleated hypopygium, thick and strong ovipositor (with basal width 3–5× its apical width posterior to constriction), antenna shorter than body, and maximum height of mesoscutellum lunules 0.4 × maximum height of lateral face of mesoscutellum. The group is strongly supported by the Bayesian molecular analysis (PP: 1.0, Fig. 1). Hosts: Pyralidae. The described species are from ACG.

Key to species of the glenriverai group
1 Metatibia almost entirely dark brown, with at most anterior 0.2 yellow; body length at most 2.3 mm and fore wing length at most 2.5 mm; T1 mostly smooth, with some sculpture near postero-lateral margins (Fig. 121h); T2 mostly smooth (Fig. 121h); fore wing with vein r 2.3 × vein 2RS; ocular-ocellar line 2.6 × posterior ocellus diameter; interocellar distance 2.1 × posterior ocellus diameter; flagellomerus 2 2.7 × as long as wide Apanteles pablovasquezi Fernández-Triana, sp. n.
Metatibia with anterior 0.5–0.6 yellow; body length at least 2.7 mm and fore wing length at least 2.8 mm; T1 with strong longitudinally striate sculpture at least on posterior 0.5 (Fig. 120f); T2 with some sculpture near posterior margin (Fig. 120f); fore wing with vein r 1.8 × vein 2RS; ocular-ocellar line 2.3 × posterior ocellus diameter; interocellar distance 1.9 × posterior ocellus diameter; flagellomerus 2 2.4 × as long as wide Apanteles glenriverai Fernández-Triana, sp. n.
guadaluperodriguezae species-group

This is a somewhat artificial group, not supported by molecular data, although the two component species share some morphological resemblance as well as similar hosts. It also looks morphologically similar to the diatraeae species-group, sharing with the latter a somewhat depressed body (dorso-ventrally), short antenna, and relatively small body size. However, it does not have the elongate and parallel-sided propodeal areola that Austin and Dangerfield (1989) considered as an apomorphic trait defining the diatraeae species-group. Additionally, the guadaluperodriguezae group, unlike the diatraeae group, attacks leaf-rolling Crambidae. The described species are from ACG.

Key to species of the guadaluperodriguezae group
1 T1 at least 4.5 × as long as its posterior width (Fig. 122f); fore wing with veins r and 2RS meeting in a smooth angle, vein 3RSa absent (Fig. 122b) [Hosts: Piletosoma thialis. A total of 30 diagnostic characters in the barcoding region: 81 C, 86 A, 88 T, 91 G, 133 A, 172 T, 250 C, 274 A, 277 T, 310 C, 313 A, 325 A, 328 T, 359 C, 361 T, 364 A, 367 C, 400 A, 412 T, 418 T, 421 A, 424 C, 472 A, 500 T, 517 A, 529 C, 595 C, 631 T, 646 T, 658 C] Apanteles guadaluperodriguezae Fernández-Triana, sp. n.
T1 at most 2.8 × as long as its posterior width (Fig. 123f); fore wing with veins r and 2RS meeting in a strong angle from where a clear vein 3RSa is visible (sometimes as a stub) (Fig. 123b) [Hosts: Pantographa expansalis, Phostria mapetalis. A total of 30 diagnostic characters in the barcoding region: 67 T, 91 T, 92 C, 136 C, 205 C, 212 C, 214 T, 217 A, 223 A, 235 T, 274 C, 299 G, 304 C, 313 C, 370 T, 379 C, 389 G, 391 T, 400 T, 421 C, 424 T, 433 T, 442 C, 481 C, 484 C, 499 T, 505 C, 542 C, 547 T, 548 C, 550 T, 565 T, 574 A, 604 C, 616 T, 622 A] Apanteles marcobustosi Fernández-Triana, sp. n.
humbertolopezi species-group

This group, comprising two species, should only be considered as interim, based on morphological evidence (strong, longitudinally striate sculpture on mediotergite 1; mediotergite 2 fully sculptured; all coxae black; pterostigma and most of veins on fore wing brown), although it is not supported by molecular data. Hosts: Elachistidae. All described species are from ACG, although we have seen other Neotropical species with similarly strong sculpture on mediotergites 1 and 2.

Key to species of the humbertolopezi group
1 Ovipositor sheaths 0.9 × as long as metatibia (Fig. 125a, c); pterostigma brown with pale spot at base (Fig. 125b); body length 2.2 mm; fore wing length 2.3 mm; flagellomerus 2 2.7 as long as wide Apanteles humbertolopezi Fernández-Triana, sp. n. (N = 1)
Ovipositor sheaths 1.2 × as long as metatibia (Fig. 126a, c); pterostigma brown (Fig. 126b); body length 2.6 mm; fore wing length 2.6 mm; flagellomerus 2 3.2 as long as wide Apanteles pablotranai Fernández-Triana, sp. n. (N = 1)
isidrochaconi species-group

This group comprises two species, characterized by extensive yellow coloration, smooth mediotergite 2, and ovipositor sheaths 1.4 × as long as metatibia. The long ovipositor differentiates this group from the rest of the Mesoamerican species with extensive yellow coloration (which usually have ovipositor sheaths shorter than metatibia, at most 1.2 × as long in a few cases). Also, the barcode for isidrochaconi is relatively unique (there is no molecular data for the other species) and provide additional support to consider this as a group on its own. There are no host records known – both species were collected by Malaise traps. Further study on its biology and/or additional DNA data will help to clarify the limits of this group in the future. The described species are from ACG.

Key to species of the isidrochaconi species-group
1 T3, laterotergites 1–3, sternites, and hypopygium mostly yellow (at most light brown near margins of T3 and hypopygium) (Figs 127a, c, f); fore and middle legs, and metacoxa entirely orange-yellow (Figs 127a, e); mesoscutellum with maximum height of lunules 0.5 × maximum height of lateral face of mesoscutellum (Fig. 127f) Apanteles isidrochaconi Fernández-Triana, sp. n. (N = 1)
T3 completely, and most of laterotergites 1–3, sternites, and hypopygium dark brown to black (Figs 128a, c, f); fore and middle legs yellow-white, metacoxa yellow-white except for anterior 0.1 which is dark brown (Fig. 128a); mesoscutellum with maximum height of lunules 0.2–0.3 × maximum height of lateral face of mesoscutellum (Fig. 128f) Apanteles juanapui Fernández-Triana, sp. n. (N = 1)
javierobandoi species-group

This comprises two species, characterized by glossa elongate (Figs 130e, 131e), tegula and humeral complex of same color (dark brown), and ovipositor about the same width from base to apex. Although the molecular data does not support the grouping of these species, and host information is only available for one of them, we have decided to consider them as a group because the combination of morphological characters detailed above is unique among Mesoamerican Apanteles. However, this group should be considered as preliminary and further study may change its status in the future. Hosts: Choreutidae. All described species are from ACG.

Key to species of the javierobandoi group
1 Antenna shorter than body, at most extending to half of metasoma; body length and fore wing length 2.4 mm; T1 length 2.4 × its width at posterior margin; T2 mostly sculptured Apanteles juangazoi Fernández-Triana, sp. n. (N = 1)
Antenna about same length or slightly larger than body; body length 2.5–3.0 mm, and fore wing length 2.6–3.0 mm; T1 length at most 2.0 × its width at posterior margin; T2 mostly smooth Apanteles javierobandoi Fernández-Triana, sp. n. (N = 4)
joserasi species-group

This group comprises one described species, although we have seen another undescribed species from the same area (with the interim name Apanteles Rodriguez79) which is only known from a male in poor condition and cannot be described in this paper. It is characterized by glossa elongate; ovipositor relatively thick and strong (with basal width more than 3.0 × its apical width posterior to constriction); maximum height of mesoscutellum lunules 0.7 × maximum height of lateral face of mesoscutellum; and propodeum with strong sculpture limited to anterior half, with posterior half mostly smooth and shiny, and with transverse carinae complete and strongly raised. All morphological traits mentioned above are similar to the leucostigmus species-group, and it might be that in the future this group is sunk within the much larger and widespread leucostigmus. However, molecular data (Fig. 1) as well as biological data (species are solitary and parasitize Venada in the joserasi group, whereas all known species in the leucostigmus group are gregarious and parasitize many genera of Eudaminae but not Venada) suggest that joserasi is better considered as a disctinct group for the time being. Hosts: Hesperiidae. The described species is from ACG.

keineraragoni species-group

This group includes two species, characterized by ovipositor sheaths half the length of metatibia, relatively short inner metatibial spur (at most 0.4 × as long as first segment of metatarsus), and body extensively dark brown to black (including full meso- and meta- soma, and all coxae). All other known species of Mesoamerican Apanteles with relatively short ovipositor sheats (i.e., 0.6 × or shorter than metatibia) have a rather extensive yellow-orange coloration. The molecular data does not support this group (Fig. 1), nor does it biology (one species is solitary on crambids, and the other is gregarious on riodinids), but we have decided to keep it as a single group for now based on the distinctive morphological traits. Hosts: Crambidae, Riodinidae. The described species are from ACG.

Key to species of the keineraragoni group
1 Fore wing with vein r 1.4 × as long as vein 2RS, vein 2M 1.5 × as long as vein (RS+M)b; flagellomerus 2 2.7 × as long as wide; interocellar distance 1.3 × as long as posterior ocelli diameter; metatibia dark brown to black on posterior 0.8 (Figs 136a, c) [Hosts: Crambidae] Apanteles keineraragoni Fernández-Triana, sp. n. (N = 3)
Fore wing with vein r 1.7 × as long as vein 2RS, vein 2M 0.7 × as long as vein (RS+M)b; flagellomerus 2 3.2 × as long as wide; interocellar distance 1.7 × as long as posterior ocelli diameter; metatibia dark brown to black on posterior 0.4–0.5 (Figs 137a, c) [Hosts: Riodinidae] Apanteles ronaldnavarroi Fernández-Triana, sp. n. (N = 1)
leucostigmus species-group

This group, by far the largest in Mesoamerica, comprises 39 species in this paper. It is defined by a thick ovipositor (as thick or thicker than the width of the median flagellomeres, and with anterior width 3.0–5.0 × its posterior width beyond the constriction), ovipositor sheaths 0.5–1.1 × as long as metatibia, propodeum with strong sculpture limited to anterior half, the posterior half mostly smooth; mesoscutellum with lateral face bearing a polished area 0.7 × or more the height of the face, pterostigma and most of fore wing white or transparent, and mediotergite 1 widening towards posterior 0.7, then narrowing toward posterior margin. The group is supported by the Bayesian molecular analysis (PP: 0.74, Fig. 1). Hosts: Hesperiidae. Widely distributed in the Neotropics; we have seen many more undescribed species in collections.

This is the only group where we extensively used molecular (i.e., barcoding) and biological (i.e., host records) characters in the key. Likewise, the species descriptions were also simplified and only include some morphological traits (plus full details on barcoding and host data). This was mostly due to the paucity of morphological characters that serve to distinguish different species. Relying solely on DNA barcoding and/or host data to describe and key species has been done before in Braconidae (e.g., Butcher et al. 2012).

However, we did some preliminary study of using morphometrics to separate species, and the results (unpublished) suggest that morphometrics may work for many, although not all, of the species in this group. We describe here the species that have been found in ACG for the sake of completing its inventory of Apanteles.

Key to species of the leucostigmus group

The species Apanteles albinervis, included in this group because of its morphology, is only known from the male holotype, and our key is only to females. There are no hosts or molecular data available for the holotype, collected in “Mexico” in 1904. It is therefore impossible to key this species by any of the character systems used here.

1 Metatibia entirely or mostly (>0.7) dark brown to black, with yellow to white usually restricted to anterior 0.2 at most (rarely with pale area extending up to anterior 0.3 of metatibia) (as in Figs 166a, d) 2
Metatibia light yellow to orange-yellow from 0.4 to almost entire metatibia (as in Figs 197c, 200c) 26
2(1) Ovipositor sheaths at least 1.0 × as long as metatibia and 1.3 × as long as metafemur 3
Ovipositor sheaths at most 0.9 × as long as metatibia and 1.1 × as long as metafemur 4
3(2) T1 length 2.7–2.8 × its width at posterior margin; T1 maximum width 1.6–1.7 × its width at posterior margin; metafemur usually more than 3.0 × as long as wide (rarely 2.8–2.9 ×) [Host species Codatractus imalena] Apanteles luzmariaromeroae Fernández-Triana, sp. n.
T1 length 2.5–2.6 × its width at posterior margin; T1 maximum width 1.4–1.5 × its width at posterior margin; metafemur 2.8 × as long as wide [Host species Astraptus talus] Apanteles marcovenicioi Fernández-Triana, sp. n. (N = 1)
4(2) Ovipositor at most 0.7 × as long as metatibia and 0.8 × as long as metafemur 5
Ovipositor more than 0.7 × as long as metatibia and usually more than 0.8 × as long as metafemur 6
5(4) Larger species, body length usually 2.3-2.5 mm (rarely 2.1 mm), and fore wing length usually 2.5–2.6 mm (rarely 2.3–2.4 mm); T1 length 2.7–2.8 × its width at posterior margin [Host species: Bungalotis erythus] Apanteles ciriloumanai Fernández-Triana, sp. n.
Smaller species, body length at most 2.1 mm, and fore wing length at most 2.3 mm; T1 length 2.5-2.6 × its width at posterior margin [Host species: Nascus spp.] Apanteles josecortesi Fernández-Triana, sp. n.
6(4) Metafemur at most 2.8 × as long as wide (rarely 2.9 × in individual specimens), and ovipositor sheaths less than 0.9 × as long as metafemur 7
Metafemur at least 2.9 × as long as wide and/or ovipositor sheaths at least 0.9 × as long as metafemur 9
7(6) Fore wing length 2.5–2.6 mm and body length at least 2.3 mm (usually more) [Host species: Ocyba calathana. A total of 18 diagnostic characters in the barcoding region: 38 C, 55 C, 61 C, 154 C, 235 T, 310 C, 316 T, 322 T, 358 C, 397 C, 405 G, 431 C, 457 C, 476 C, 604 T, 610 C, 637 A, 641 C] Apanteles cynthiacorderoae Fernández-Triana, sp. n.
Fore wing length at most 2.4 mm (usually less) and body length usually less than 2.3 mm [Host species: Cephise aelius or Phocides spp. A total of 18 diagnostic characters in the barcoding region: 38 T, 55 T, 61 T, 154 T, 235 C, 310 T, 316 A, 322 A, 358 T, 397 T, 405 A, 431 A, 457 T, 476 A, 604 A, 610 T, 637 T, 641 T] 8
8(7) T1 length 2.3–2.8 × its width at posterior margin (rarely 2.1–2.2 ×) [Host species: Cephise aelius. A total of 39 diagnostic characters in the barcoding region: 19 T, 43 A, 49 C, 98 A, 118 C, 170 A, 181 G, 184 A, 187 T, 212 C, 238 T, 259 C, 263 T, 284 C, 295 A, 298 A, 304 T, 340 C, 364 T, 379 T, 400 C, 421 T, 439 C, 448 T, 458 T, 490 C, 507 T, 508 T, 529 C, 536 T, 562 A, 574 A, 578 T, 589 T, 601 C, 616 T, 629 T, 646 T, 652 C] Apanteles hazelcambroneroae Fernández-Triana, sp. n.
T1 length 2.1–2.2 × its width at posterior margin [Host species: Phocides spp. A total of 39 diagnostic characters in the barcoding region: 19 C, 43 T, 49 T, 98 G, 118 T, 170 G, 181 A, 184 T, 187 C, 212 T, 238 C, 259 T, 263 C, 284 T, 295 T, 298 G, 304 C, 340 T, 364 A, 379 C, 400 T, 421 C, 439 T, 448 C, 458 C, 490 T, 507 C, 508 C, 529 T, 536 C, 562 T, 574 T, 578 C, 589 C, 601 T, 616 C, 629 C, 646 C, 652 T] Apanteles randallgarciai Fernández-Triana, sp. n.
9(6) Fore wing with veins C+Sc+R and R1 mostly brown; usually veins r, 2RS, 2M, (RS+M)b, 1CU, 2Cua, and 1m-cu partially brown; interior area of other veins, and at least part of pterostigma, usually light brown or yellowish-white (as in Figs 165b, 172b, 189b) 10
Fore wing with veins C+Sc+R and R1 with brown coloration restricted narrowly to borders, interior area of those veins and pterostigma (and sometimes veins r, 2RS and 2M) transparent or white; other veins mostly transparent (as in Figs 173b, 174b, 175b) 19
10(9) Metafemur 2.7 × as long as wide; ovipositor sheaths 0.9 × as long as metatibia and 1.1 × as long as metafemur Apanteles eugeniaphilipsae Fernández-Triana, sp. n. (N = 2)
Metafemur at least 2.8 × as long as wide; ovipositor sheaths at most 0.8 × (rarely 0.9 ×) as long as metatibia and at most 1.0 × as long as metafemur 11
11(10) Maximum width of T1 (at about 0.7–0.8 × its length) more than 1.7 × its width at posterior margin Apanteles rodrigogamezi Fernández-Triana, sp. n.
Maximum width of T1 (at about 0.7–0.8 × its length) less than 1.6 × its width at posterior margin 12
12(11) Maximum width of T1 (at about 0.7–0.8 × its length) usually at most 1.2 × its width at posterior margin; T1 appearing almost parallel-sided Apanteles gerardobandoi Fernández-Triana, sp. n.
Maximum width of T1 at least 1.3 × its width at posterior margin; T1 clearly appearing to widen from base to 0.7–0.8 × its length, then narrowing towards posterior margin of mediotergite 13
13(12) Ovipositor sheaths about 0.44 mm, metafemur 0.47 mm, metatibia 0.59 mm, and maximum width of T1 0.18 mm, much shorter than below; body length 1.9–2.0 mm and fore wing 2.1–2.2 mm Apanteles ricardocaleroi Fernández-Triana, sp. n.
Ovipositor sheaths 0.49–0.59 mm, metafemur 0.54–0.59 mm, metatibia 0.63–0.72 mm and maximum width of T1 0.20–0.25 mm, much longer than above; body length and fore wing usually larger than 2.2 mm, very rarely smaller 14
14(13) Ovipositor sheaths at most 2.0 × (rarely 2.3 ×) as long as maximum width of T1 Apanteles diniamartinezae Fernández-Triana, sp. n.
Ovipositor sheaths at least 2.4 × as long as maximum width of T1 15
15(14) Host species: Calliades zeutus or Urbanus doryssus 16
Hosts species: Telemiades spp. (one single rearing record from Phocides lilea) 17
16(15) Body length 1.9–2.0 mm; fore wing 2.1–2.2 mm [Host species: Calliades zeutus. A total of 23 diagnostic characters in the barcoding region: 30 C, 66 G, 75 G, 84 T, 138 T, 147 A, 192 T, 219 T, 264 A, 315 A, 352 C, 378 T, 388 A, 397 T, 414 A, 420 C, 528 C, 535 T, 547 T, 561 T, 627 T, 639 C, 645 C] Apanteles pabloumanai Fernández-Triana, sp. n.
Body length 2.3 mm or more (rarely 2.1 mm); fore wing at least 2.5 mm [Host species: Urbanus doryssus. A total of 23 diagnostic characters in the barcoding region: 30 T, 66 A, 75 A, 84 C, 138 C, 147 G, 192 C, 219 C, 264 G, 315 T, 352 T, 378 C, 388 G, 397 G, 414 G, 420 A, 528 T, 535 C, 547 C, 561 A, 627 A, 639 T, 645 T] Apanteles josemonteroi Fernández-Triana, sp. n.
17(15) Host species: Telemiades oiclus. A total of 10 diagnostic characters in the barcoding region: 57 G, 144 T, 264 G, 273 C, 276 T, 339 C, 381 G, 477 T, 525 C, 645 C Apanteles carlosviquezi Fernández-Triana, sp. n.
Hosts species: Telemiades fides (one single rearing record from Phocides lilea). A total of 10 diagnostic characters in the barcoding region: 57 A, 144 C, 264 A, 273 T, 276 A, 339 T, 381 A, 477 A, 525 T, 645 T 18
18(17) A total of 18 diagnostic characters in the barcoding region: 73 C, 99 A, 205 C, 265 T, 270 T, 286 C, 315 T, 321 A, 358 T, 462 C, 489 T, 528 T, 535 T, 541 T, 564 T, 567 T, 573 A, 624 A, Apanteles inesolisae Fernández-Triana, sp. n.
A total of 18 diagnostic characters in the barcoding region: 73 T, 99 G, 205 T, 265 C, 270 C, 286 T, 315 A, 312 T, 358 C, 462 T, 489 C, 528 C, 535 C, 541 C, 564 A, 567 C, 573 C, 624 T Apanteles manuelzumbadoi Fernández-Triana, sp. n.
19(9) Ovipositor sheaths 0.6–0.8 × (average 0.7 ×) as long as metatibia and 0.8–0.9 × as long as metafemur 20
Ovipositor sheaths 0.8–0.9 × (average at least 0.8 ×) as long as metatibia and at least 1.0 × as long as metafemur 21
20(19) Antenna same length or longer than body; T1 length usually less than 2.3 × its width at posterior margin; ovipositor sheaths 0.7–0.8 × as long as metatibia and 0.8–1.0 × as long as metafemur Apanteles raulsolorsanoi Fernández-Triana, sp. n.
Antenna shorter than body; T1 length 2.5–2.6 × its width at posterior margin; ovipositor sheaths 0.5–0.6 × as long as metatibia and 0.7–0.8 × as long as metafemur Apanteles juanmatai Fernández-Triana, sp. n.
21(19) Host species: Aguna spp 22
Host species: either Bungalotis, Chioides, Polygonus, Telemiades, or Urbanus 23
22(21) Metatibia almost entirely dark brown to black, with yellow to white coloration restricted to anterior 0.1 at most; T1 length 2.3–2.4 × its width at posterior margin; T1 maximum width 1.2–1.3 × its width at posterior margin Apanteles minorcarmonai Fernández-Triana, sp. n.
Metatibia with anterior 0.3 yellow; T1 length 2.9 × or more its width at posterior margin; T1 maximum width 1.8–1.9 × its width at posterior margin Apanteles jesusugaldei Fernández-Triana, sp. n.
23(21) Antenna clearly shorter than body length, usually 0.8–0.9 × as long as body; metatibia with anterior 0.3 yellow (a few specimens may have metatibia anterior 0.5 yellow, and will not run through here) 24
Antenna as long or slightly longer than body length; metatibia almost entirely dark brown to black, with yellow to white coloration restricted to anterior 0.1 at most 25
24(23) T1 length more than 3.0 × its width at posterior margin; T1 maximum width 1.8–1.9 × its width at posterior margin [Host species: Urbanus spp.] Apanteles eliethcantillanoae Fernández-Triana, sp. n.
T1 length 2.3–2.4 × its width at posterior margin; T1 maximum width 1.4–1.5 × its width at posterior margin [Hosts species: Chioides zilpa, Polygonus leo] Apanteles federicomatarritai Fernández-Triana, sp. n.
25(23) Body length 2.3–2.6 mm (rarely 2.1–2.2 mm); fore wing length at least 2.5 mm; metafemur length 2.7–3.0 × its width [Host species: Bungalotis quadratum] Apanteles alvarougaldei Fernández-Triana, sp. n.
Body length 2.1–2.2 mm); fore wing length 2.3–2.4 mm; metafemur length 3.2–3.3 × its width [Host species: Telemiades fides] Apanteles johanvargasi Fernández-Triana, sp. n. (N =3)
26(2) Metatibia almost entirely yellow, at most with posterior 0.1 brown or just with slightly darker spot which is almost same color than rest of metatibia 27
Metatibia with posterior 0.3–0.4 dark brown, clearly darker than rest of metatibia 31
27(26) Ovipositor sheaths averaging 0.44 mm (range 0.40–0.46 mm), their length 0.6–0.7 × metatibia length and 0.7–0.8 × metafemur length Apanteles mariachavarriae Fernández-Triana, sp. n.
Ovipositor sheaths usually over 0.50 mm (if rarely 0.45 mm in length, then species average over 0.48 mm), ovipositor sheaths 0.8 × metatibia length (rarely 0.7 ×) and 0.9–1.0 × as long as metafemur 28
28(27) Antenna shorter than body; T1 length 2.7–2.8 × its width at posterior margin; T1 maximum width 1.6–1.7 × its width at posterior margin Apanteles duvalierbricenoi Fernández-Triana, sp. n.
Antenna at least as long as body; T1 length 2.3-2.4 × its width at posterior margin; T1 maximum width 1.4–1.5 × its width at posterior margin 29
29(28) Host species: Astraptes anaphus. A total of 14 diagnostic characters in the barcoding region: 73 T, 145 C, 193 T, 265 A, 293 A, 316 A, 343 G, 359 C, 401 C, 421 T, 476 C, 562 T, 571 C, 628 T Apanteles sigifredomarini Fernández-Triana, sp. n.
Host species: Urbanus spp. (in two rare cases Astraptes alardus, Dan check that for species Rodriguez24). A total of 14 diagnostic characters in the barcoding region: 73 C, 145 T, 193 C, 265 G, 293 T, 316 T, 343 A, 359 T, 401 T, 421 A, 476 A, 562 A, 571 T, 628 A 30
30(29) Host species: Urbanus simplicius. A total of four diagnostic characters in the barcoding region: 166 G, 232 C, 373 T, 379 T Apanteles sergioriosi Fernández-Triana, sp. n.
Host species: Urbanus dorantes (plus 2 Astraptes records). A total of four diagnostic characters in the barcoding region: 166 A, 232 A, 373 A, 379 C Apanteles ronaldzunigai Fernández-Triana, sp. n.
31(26) Fore wing with veins C+Sc+R and R1 mostly brown; usually veins r, 2RS, 2M, (RS+M)b, 1CU, 2Cua, and 1m-cu partially brown; interior area of other veins, and at least part of pterostigma, usually light brown or yellowish-white (as in Figs 165b, 172b, 189b) 32
Fore wing with veins C+Sc+R and R1 with brown coloration restricted narrowly to borders, interior area of those veins and pterostigma (and sometimes veins r, 2RS and 2M) transparent or white; other veins mostly transparent (as in Figs 173b, 174b, 175b) 33
32(31) Ovipositor sheaths 0.8 × as long as metatibia and 1.0 × as long as metafemur; T1 length 2.7–2.8 × its width at posterior margin [Host species: Urbanus doryssus] Apanteles lilliammenae Fernández-Triana, sp. n.
Ovipositor sheaths 0.5 × as long as metatibia and 0.6 × as long as metafemur; T1 length 2.3–2.4 × its width at posterior margin [Host species: Urbanus dorantes, Urbanus teleus] Apanteles wadyobandoi Fernández-Triana, sp. n.
33(31) Ovipositor sheaths 0.7 × as long as metatibia and 0.7–0.8 × as long as metafemur; metafemur 3.2 × as long as wide 34
Ovipositor sheaths usually 0.8 × as long as metatibia (rarely 0.7 ×) and 0.9–1.0 × as long as metafemur; metafemur usually less than 3.0 × as long as wide (rarely up to 3.2 ×) 35
34(33) Body length at most 2.2 mm and fore wing length at most 2.4 mm; metafemur at most 2.9 × as long as wide; T1 length less than 2.0 × its width at posterior margin [Host species: Urbanus proteus. Distribution: Caribbean islands (Cuba, Grenada, Puerto Rico, St. Vincent), and southern United States (Florida)] Apanteles leucostigmus (Ashmead, 1900)
Body length at least 2.5 mm and fore wing length at least 2.7 mm; metafemur at least 3.2 × as long as wide; T1 length more than 2.6 × its width at posterior margin [Host species: mostly Astraptes spp., four known records of Urbanus spp. (all different species than Urbanus proteus). Distribution: Costa Rica (ACG)] Apanteles jorgehernandezi Fernández-Triana, sp. n.
35(33) T1 length 1.9–2.0 × its width at posterior margin [Host species: Mostly Urbanus albimargo and Urbanus doryssus (rarely also Autochton sp.). A total of 19 diagnostic characters in the barcoding region: 54 C, 99 A, 177 C, 186 C, 216 T, 237 T, 330 T, 343 A, 388 C, 387 T, 396 A, 423 T, 460 A, 461 T, 528 T, 534 T, 558 A, 580 T, 606 G] Apanteles rostermoragai Fernández-Triana, sp. n.
T1 length 2.3–2.6 × its width at posterior margin [Host species: Mostly Achalarus, Astraptus, Cogia and Thessia; if from genus Urbanus, then almost always from other species than above (Urbanus belli, Urbanus dorantes, Urbanus teleus and Urbanus viterboana; very rarely from Urbanus albimargo). Barcoding region with different nucleotides at positions mentioned in first half of couplet] 36
36(35) T1 length 2.5–2.6 × its width at posterior margin; T1 maximum width 1.6–1.7 × its width at posterior margin [Host species: Urbanus albimargo, and rarely from Achalarus toxeus, Cogia calchas and Thessia jalapus. A total of 10 diagnostic characters in the barcoding region: 57 C, 93 C, 111 T, 117 G, 150 T, 177 A, 183 T, 309 A, 4444 T, 606 C] Apanteles angelsolisi Fernández-Triana, sp. n.
T1 length 2.3–2.4 × its width at posterior margin; T1 maximum width 1.4–1.5 × its width at posterior margin [Host species: Astraptes spp., and Urbanus spp. but not Urbanus albimargo. Barcoding region with different nucleotides at positions mentioned in first half of couplet] 37
37(36) Metafemur length usually less than 3.0 × its width (range: 2.8–3.1 ×); fore wing length 2.2–2.5 mm [Host species: Urbanus belli (with one record of Urbanus viterboana). A total of five diagnostic characters in the barcoding region: 192 G, 225 T, 279 C, 615 C, 685 T] Apanteles gladysrojasae Fernández-Triana, sp. n.
Metafemur length usually more than 3.0 × its width (range: 3.0–3.4 ×); fore wing length 2.5–2.7 mm [Host species: Mostly species of Astraptes (Astraptes alardus, Apanteles apastus, Astraptes brevicauda, Astraptes talus, Astraptes tucuti), with one record of Urbanus belli. Barcoding region with different nucleotides at positions mentioned in first half of couplet] Apanteles bernardoespinozai Fernández-Triana, sp. n.
marisolnavarroae species-group

This group comprises two species, characterized by relatively large body size (body and fore wing length at least 3.3 mm, usually longer), mesoscutellar disc punctured, tegula and humeral complex of different color, and brown pterostigma. The group is strongly supported by the Bayesian molecular analysis (PP: 1.0, Fig. 1). Hosts: Pyralidae. All described species are from ACG.

Key to species of the marisolnavarroae group
1 Meso- and metatrochantellus yellow (Fig. 145a); metatibia mostly yellow, with only dark spot on posterior 0.1–0.2 Apanteles randallmartinezi Fernández-Triana, sp. n. (N = 2)
Meso- and metatrochantellus dark brown to black (Fig. 144a); metatibia with posterior 0.3–0.4 dark brown to black (Fig. 144c) Apanteles marisolnavarroae Fernández-Triana, sp. n. (N = 2)
megathymi species-group

This group comprises two species, characterized by the combination of relatively long ovipositor sheaths, 1.4–1.5 × as long as metatibia; mesoscutellar disc smooth, contrasting with strongly punctured anteromesoscutum; propodeum strongly carinated and sculptured; pterostigma mostly transparent, with thin brown borders; fore wing with shape of junction of veins r and 2RS strongly angulated, and often with a knob; metafemur and metatibia completely or at least partially yellow-orange; and all coxae dark brown to black. We have tentatively considered this as a group based on the morphological similarities; however, there is no molecular data available for those two species, and the host families are different. Future study might find this group to be completely artificial. Hosts: Gelechiidae, Hesperiidae. The two species are widely distributed in the New World, one mostly in the Nearctic, the other in the Neotropics.

Key to species of the megathymi group
1 Body length at least 3.5 mm, and fore wing length at least 3.7 mm; T1 length 2.4–2.8 × its posterior width; T2 mostly smooth [Hosts: Hesperiidae. Distribution: Mexico, United States] Apanteles megathymi Riley, 1881
Body length at most 3.0 mm, and fore wing length at most 3.2 mm; T1 length 1.3 × its posterior width; T2 entirely sculptured with longitudinal striation (Fig. 146f) [Hosts: Gelechiidae. Distribution: Brazil, Cuba, Grenada, St. Vincent] Apanteles balthazari (Ashmead, 1900)
paranthrenidis species-group

This group comprises four species, characterized by a relatively broad mediotergite 1 (its length at most 1.3 × its width); pterostigma transparent or whitish with only thin brown borders, and most of the fore wing veins transparent; vein 2M at most 0.6 × as long as vein (RS+M)b; and lateral face of scutellum with polished area 0.7–0.8 × maximum face height. Only for one species there are barcodes available, therefore more data will be needed for molecular analysis of this group, which is considered here as just a interim arrangement of species. Hosts: Crambidae, Gelechiidae, Noctuidae, Pyralidae, Sesiidae (some of those records may be questionable, especially those from old references). Most of the available host records are from miners. Distribution: Widely distributed in the New World.

Key to species of the paranthrenidis group
1 Femora mostly yellow-orange, at most with small dark spot on posterior 0.1–0.2 of metafemur (Figs 151a, c, 152a, c) 2
Mesofemur dark brown to black on at least anterior 0.5, metafemur entirely dark brown to reddish (Figs 150a, c, 153a, c) 3
2(1) Darker species, with all coxae dark brown to black, metafemur and metatibia with dark spot on posterior 0.1–0.2; flagellomerus 2 3.1 × as long as wide; scutellar suture with up to 13 pits; T2 mostly smooth and width at apex 3.1 × its length (Fig. 151g); fore wing with vein r 1.6 × as long as vein 2RS, and vein 2RS 1.7 × as long as vein 2M [Hosts: Crambidae] Apanteles megastidis Muesebeck, 1958
Lighter species, with at least pro- and meso- coxae light brown to yellow, metafemur and metatibia completely yellow to orange; flagellomerus 2 2.2 × as long as wide; scutellar suture with at most 10 pits; T2 with some sculpture near posterior margin and width at apex at least 3.6 × its length (usually more) (Fig. 152f); fore wing with vein r 3.0 × as long as vein 2RS, and vein 2RS 1.1 × as long as vein 2M [Hosts: Noctuidae, Sesiidae] Apanteles paranthrenidis Muesebeck, 1921
3(2) Glossa weakly elongate (Fig. 150f); tarsal claws with a basal spine-like seta; metatibia with posterior 0.3 dark; metatarsus with segment 1 dark brown to black on posterior 0.8–0.9 (Fig. 150a); interocellar distance 1.6 × as long as posterior ocellus diameter; T2 with at posterior margin 4.0 × its length; metatibial inner spur 1.8 × as long as outer spur [Hosts: Crambidae, Pyralidae] Apanteles esthercentenoae Fernández-Triana, sp. n.
Glossa not elongate (Fig. 153e); tarsal claws simple; metatibia with posterior 0.1 dark; metatarsus with segment 1 dark brown on posterior 0.5 (Fig. 153a); interocellar distance 1.9 × as long as posterior ocellus diameter; T2 with at posterior margin 3.4 × its length; metatibial inner spur 1.4 × as long as outer spur [Hosts: Gelechiidae, Noctuidae] Apanteles thurberiae Muesebeck, 1921
ronaldgutierrezi species-group

This group comprises two species, characterized by the combination of pale tegula and humeral complex dark, ovipositor sheaths much shorter (0.6 ×) than metatibia length, and metatrochanter, metatrochantellus, and anterior third of metafemur yellow-white. Molecular data also supports the species as divergent (Fig. 1). We have included here the species Apanteles insularis Muesebeck, 1921, based on the examination of few photos from the holotype (see more comments below under that species). However, we cannot be sure of the actual placement of insularis until more specimens can be examined, thus its placement here is preliminary and likely to change over time. Hosts: Choreutidae. The described species are from Costa Rica (ACG), Grenada and St. Vincent.

samarshalli species-group

This group comprises two unique species among all described Apanteles in Mesoamerica, characterized by having fore wing vein 2M very short and close to vein 2RS (almost obliterating the space of the second submarginal cell), and antenna very short, not surpassing the mesosoma. The group is strongly supported by the Bayesian molecular analysis (PP: 1.0, Fig. 1), although it clusters apart from all other known species of Mesoamerican Apanteles – strongly suggesting it might better be placed on a different genus on its own when future studies on Microgastrinae phylogeny are done. No host is known. One of the species is rather widespread (Neartic and Neotropical) whereas the other one is only know from ACG.

Key to species of the samarshalli group
1 Scape, pedicel (partially), anterior 0.7 of metatibia and anterior 0.6 of first segment of metatarsus yellow (Figs 205a, c); propodeum with areola weakly defined by central impression and few rugae or minute carinae arising from nucha (Fig. 205d) Apanteles samarshalli Fernández-Triana, 2010
Scape and pedicel light brown (Fig. 160f); posterior 0.5 of metatibia and first segment of metatarsus dark brown (Figs 160a, d); propodeum with areola completely defined by carinae (Fig. 160g) Apanteles jimmychevezi Fernández-Triana, sp. n. (N = 1)
Taxonomic treatment of species (in alphabetical order)
Type locality.

COSTA RICA, Alajuela, ACG, Sector Rincon Rain Forest, Estación Llanura, 135m, 10.93332, -85.25331.

Holotype.

♀ in CNC. Specimen labels: 1. Voucher: D.H.Janzen & W.Hallwachs, DB: http://janzen.sas.upenn.edu, Area de Conservación Guanacaste, COSTA RICA, 09-SRNP-75013. 2. DHJPAR0039774.

Description.

Female. Body color: body mostly dark except for some sternites which may be pale. Antenna color: scape, pedicel, and flagellum dark. Coxae color (pro-, meso-, metacoxa): dark, dark, dark. Femora color (pro-, meso-, metafemur): anteriorly dark/posteriorly pale, dark, dark. Tibiae color (pro-, meso-, metatibia): pale, pale, mostly dark but anterior 0.2 or less pale. Tegula and humeral complex color: tegula pale, humeral complex half pale/half dark. Pterostigma color: mostly pale and/or transparent, with thin dark borders. Fore wing veins color: partially pigmented (a few veins may be dark but most are pale). Antenna length/body length: antenna about as long as body (head to apex of metasoma); if slightly shorter, at least extending beyond anterior 0.7 metasoma length. Body in lateral view: not distinctly flattened dorso–ventrally. Body length (head to apex of metasoma): 3.1–3.2 mm. Fore wing length: 3.1–3.2 mm. Ocular–ocellar line/posterior ocellus diameter: 2.3–2.5. Interocellar distance/posterior ocellus diameter: 1.7–1.9. Antennal flagellomerus 2 length/width: 2.9–3.1. Antennal flagellomerus 14 length/width: 1.4–1.6. Length of flagellomerus 2/length of flagellomerus 14: 2.0–2.2. Tarsal claws: simple. Metafemur length/width: 3.2–3.3. Metatibia inner spur length/metabasitarsus length: 0.4–0.5. Anteromesoscutum: mostly with deep, dense punctures (separated by less than 2.0 × its maximum diameter). Mesoscutellar disc: mostly smooth. Number of pits in scutoscutellar sulcus: 9 or 10. Maximum height of mesoscutellum lunules/maximum height of lateral face of mesoscutellum: 0.6–0.7. Propodeum areola: completely defined by carinae, including transverse carina extending to spiracle. Propodeum background sculpture: mostly sculptured. Mediotergite 1 length/width at posterior margin: 2.6–2.8. Mediotergite 1 shape: mostly parallel–sided for 0.5–0.7 of its length, then narrowing posteriorly so mediotergite anterior width >1.1 × posterior width. Mediotergite 1 sculpture: mostly sculptured, excavated area centrally with transverse striation inside and/or a polished knob centrally on posterior margin of mediotergite. Mediotergite 2 width at posterior margin/length: 2.8–3.1. Mediotergite 2 sculpture: mostly smooth. Outer margin of hypopygium: with a wide, medially folded, transparent, semi–desclerotized area; usually with 4 or more pleats. Ovipositor thickness: about same width throughout its length. Ovipositor sheaths length/metatibial length: 1.2–1.3. Length of fore wing veins r/2RS: 1.7–1.9. Length of fore wing veins 2RS/2M: 2.1 or more. Length of fore wing veins 2M/(RS+M)b: 0.5–0.6. Pterostigma length/width: 3.1–3.5. Point of insertion of vein r in pterostigma: clearly beyond half way point length of pterostigma. Angle of vein r with fore wing anterior margin: more or less perpendicular to fore wing margin. Shape of junction of veins r and 2RS in fore wing: distinctly but not strongly angled.

Male. Unknown.

Molecular data.

Sequences in BOLD: 2, barcode compliant sequences: 2.

Biology/ecology.

Solitary (Fig. 210). Hosts: Elachistidae, Antaeotricha Janzen86, Stenoma Janzen148.

Distribution.

Costa Rica, ACG.

Etymology.

We dedicate this species to Adelina Morales for her diligent efforts as a parataxonomist in the ACG inventory of its plant viruses and for Estación Biológica Santa Rosa.

Type locality.

COSTA RICA, Alajuela, ACG, Sector Rincon Rain Forest, Sendero Tucan, 410m, 10.90424, -85.2712.

Holotype.

♀ in CNC. Specimen labels: 1. DHJPAR0039757. 2. COSTA RICA, Alajuela, ACG, Sector Rincon Rain Forest, Sendero Tucan, 10.90424, -85.2712, 410m, 16.vii.2009, DHJPAR0039757. 3. Voucher: D.H.Janzen & W.Hallwachs, DB: http://janzen.sas.upenn.edu, Area de Conservación Guanacaste, COSTA RICA, 09-SRNP-41735.

Paratypes.

2 ♀, 3 ♂ (CNC, NMNH). COSTA RICA, ACG database codes: DHJPAR0048170, DHJPAR0039073.

Description.

Female. Body color: body mostly dark except for some sternites which may be pale. Antenna color: scape, pedicel, and flagellum dark. Coxae color (pro-, meso-, metacoxa): dark, dark, dark. Femora color (pro-, meso-, metafemur): anteriorly dark/posteriorly pale, dark, dark. Tibiae color (pro-, meso-, metatibia): pale, pale, mostly dark but anterior 0.2 or less pale. Tegula and humeral complex color: tegula pale, humeral complex half pale/half dark. Pterostigma color: mostly pale and/or transparent, with thin dark borders. Fore wing veins color: partially pigmented (a few veins may be dark but most are pale). Antenna length/body length: antenna about as long as body (head to apex of metasoma); if slightly shorter, at least extending beyond anterior 0.7 metasoma length. Body in lateral view: not distinctly flattened dorso–ventrally. Body length (head to apex of metasoma): 2.5–2.6 mm. Fore wing length: 2.7–2.8 mm or 2.9–3.0 mm. Ocular–ocellar line/posterior ocellus diameter: 2.3–2.5. Interocellar distance/posterior ocellus diameter: 2.0–2.2. Antennal flagellomerus 2 length/width: 2.6–2.8. Antennal flagellomerus 14 length/width: 1.7–1.9. Length of flagellomerus 2/length of flagellomerus 14: 1.7–1.9. Tarsal claws: with single basal spine–like seta. Metafemur length/width: 3.0–3.1. Metatibia inner spur length/metabasitarsus length: 0.4–0.5. Anteromesoscutum: mostly with shallow, dense punctures (separated by less than 2.0 × its maximum diameter). Mesoscutellar disc: with a few sparse punctures. Number of pits in scutoscutellar sulcus: 9 or 10. Maximum height of mesoscutellum lunules/maximum height of lateral face of mesoscutellum: 0.8 or more. Propodeum areola: completely defined by carinae, including transverse carina extending to spiracle. Propodeum background sculpture: partly sculptured, especially on anterior 0.5. Mediotergite 1 length/width at posterior margin: 1.7–1.9. Mediotergite 1 shape: more or less parallel–sided. Mediotergite 1 sculpture: mostly sculptured, excavated area centrally with transverse striation inside and/or a polished knob centrally on posterior margin of mediotergite. Mediotergite 2 width at posterior margin/length: 3.6–3.9. Mediotergite 2 sculpture: mostly smooth. Outer margin of hypopygium: with a wide, medially folded, transparent, semi–desclerotized area; usually with 4 or more pleats. Ovipositor thickness: about same width throughout its length. Ovipositor sheaths length/metatibial length: 1.4–1.5. Length of fore wing veins r/2RS: 2.3 or more. Length of fore wing veins 2RS/2M: 0.9–1.0. Length of fore wing veins 2M/(RS+M)b: 0.5–0.6. Pterostigma length/width: 3.6 or more. Point of insertion of vein r in pterostigma: about half way point length of pterostigma. Angle of vein r with fore wing anterior margin: clearly outwards, inclined towards fore wing apex. Shape of junction of veins r and 2RS in fore wing: distinctly but not strongly angled.

Male. As in female, with slender mediotergite 1.

Molecular data.

Sequences in BOLD: 3, barcode compliant sequences: 3.

Biology/ecology.

Solitary (Fig. 225). Host: Elachistidae, Stenoma Janzen08 feeding on Clusia spp.

Distribution.

Costa Rica, ACG.

Etymology.

We dedicate this species to Adriana Chavarría in recognition of her diligent efforts for the ACG Programa de Ecoturismo.

Apanteles adrianaguilarae Fernández-Triana, sp. n.

http://zoobank.org/73C1363A-38F8-408F-9F74-5DCBC2D10AC8

http://species-id.net/wiki/Apanteles_adrianaguilarae

Figs 32, 232
Apanteles Rodriguez15. Smith et al. (2008). Interim name provided by the authors.
Type locality.

COSTA RICA, Alajuela, ACG, Sector Rincón Rain Forest, Rio Francia Arriba, 400m, 10.89666, -85.29003.

Holotype.

♀ in CNC. Specimen labels: 1. COSTA RICA, Alajuela, ACG, Sector Rincón Rain Forest, Rio Francia Arriba, 27.vii.2001, 400m, 10.89666, -85.29003, DHJPAR0001553.

Paratypes.

43 ♀, 14 ♂ (BMNH, CNC, INBIO, INHS, NMNH). COSTA RICA, ACG database codes: DHJPAR0003005, DHJPAR0003027, DHJPAR0034265, DHJPAR0034271, DHJPAR0038956, 01-SRNP-5505, 02-SRNP-1979, 04-SRNP-34656, 04-SRNP-34908, 04-SRNP-55638, 04-SRNP-55691.

Description.

Female. Body color: head dark, mesosoma dark with parts of axillar complex pale, metasoma with some mediotergites, most laterotergites, sternites, and/or hypopygium pale. Antenna color: scape, pedicel, and flagellum pale. Coxae color (pro-, meso-, metacoxa): pale, pale, pale or pale, pale, partially pale/partially dark. Femora color (pro-, meso-, metafemur): pale, pale, pale. Tibiae color (pro-, meso-, metatibia): pale, pale, mostly pale but with posterior 0.2 or less dark. Tegula and humeral complex color: both pale. Pterostigma color: dark. Fore wing veins color: mostly dark (a few veins may be unpigmented). Antenna length/body length: antenna about as long as body (head to apex of metasoma); if slightly shorter, at least extending beyond anterior 0.7 metasoma length. Body in lateral view: not distinctly flattened dorso–ventrally. Body length (head to apex of metasoma): 2.7–2.8 mm or 2.9–3.0 mm. Fore wing length: 2.7–2.8 mm, 2.9–3.0 mm or 3.1–3.2 mm. Ocular–ocellar line/posterior ocellus diameter: 2.3–2.5. Interocellar distance/posterior ocellus diameter: 2.0–2.2. Antennal flagellomerus 2 length/width: 2.6–2.8. Antennal flagellomerus 14 length/width: 1.7–1.9. Length of flagellomerus 2/length of flagellomerus 14: 2.0–2.2. Tarsal claws: simple or with single basal spine–like seta. Metafemur length/width: 2.8–2.9. Metatibia inner spur length/metabasitarsus length: 0.4–0.5. Anteromesoscutum: mostly with deep, dense punctures (separated by less than 2.0 × its maximum diameter). Mesoscutellar disc: with a few sparse punctures. Number of pits in scutoscutellar sulcus: 7 or 8. Maximum height of mesoscutellum lunules/maximum height of lateral face of mesoscutellum: 0.4–0.5. Propodeum areola: completely defined by carinae, but only partial or absent transverse carina. Propodeum background sculpture: mostly sculptured. Mediotergite 1 length/width at posterior margin: 2.3–2.5. Mediotergite 1 shape: mostly parallel–sided for 0.5–0.7 of its length, then narrowing posteriorly so mediotergite anterior width >1.1 × posterior width. Mediotergite 1 sculpture: with some sculpture near lateral margins and/or posterior 0.2–0.4 of mediotergite. Mediotergite 2 width at posterior margin/length: 4.4–4.7. Mediotergite 2 sculpture: mostly smooth. Outer margin of hypopygium: with a wide, medially folded, transparent, semi–desclerotized area; usually with 4 or more pleats. Ovipositor thickness: about same width throughout its length. Ovipositor sheaths length/metatibial length: 0.6–0.7. Length of fore wing veins r/2RS: 1.4–1.6. Length of fore wing veins 2RS/2M: 1.1–1.3. Length of fore wing veins 2M/(RS+M)b: 0.9–1.0. Pterostigma length/width: 3.1–3.5. Point of insertion of vein r in pterostigma: clearly beyond half way point length of pterostigma. Angle of vein r with fore wing anterior margin: clearly outwards, inclined towards fore wing apex. Shape of junction of veins r and 2RS in fore wing: strongly angulated, sometimes with a knob.

Male. Metacoxa tends to have an anterodorsal brown spot, otherwise similar to female.

Molecular data.

Sequences in BOLD: 37, barcode compliant sequences: 37.

Biology/ecology.

Gregarious (Fig. 232). Host: Tortricidae, Anacrusis nephrodes.

Distribution.

Costa Rica, ACG.

Etymology.

We dedicate this species to Adriana Aguilar in recogition of her diligent efforts for the ACG Programa Forestal.

Type locality.

COSTA RICA, Guanacaste, ACG, Potrerillos, Río Azufrado, 95m, 10.81224, -85.54438.

Holotype.

♀ in CNC. Specimen labels: 1. DHJPAR0005279. 2. COSTA RICA, Guanacaste, ACG, Potrerillos, Río Azufrado, 23.vii.2000, gusaneros. 3. 00-SRNP-16110, Same as 00-16047, On Inga vera.

Paratypes.

1 ♂ (CNC). COSTA RICA, ACG database codes: DHJPAR0039780).

Description.

Female. Body color: body mostly dark except for some sternites which may be pale. Antenna color: scape, pedicel, and flagellum dark. Coxae color (pro-, meso-, metacoxa): dark, dark, dark. Femora color (pro-, meso-, metafemur): anteriorly dark/posteriorly pale, dark, dark. Tibiae color (pro-, meso-, metatibia): pale, anteriorly pale/posteriorly dark, dark. Tegula and humeral complex color: tegula dark, humeral complex half pale/half dark. Pterostigma color: dark with pale spot at base. Fore wing veins color: partially pigmented (a few veins may be dark but most are pale). Antenna length/body length: antenna about as long as body (head to apex of metasoma); if slightly shorter, at least extending beyond anterior 0.7 metasoma length. Body in lateral view: not distinctly flattened dorso–ventrally. Body length (head to apex of metasoma): 2.5–2.6 mm. Fore wing length: 2.5–2.6 mm. Ocular–ocellar line/posterior ocellus diameter: 2.3–2.5. Interocellar distance/posterior ocellus diameter: 1.7–1.9. Antennal flagellomerus 2 length/width: 2.6–2.8. Antennal flagellomerus 14 length/width: 1.4–1.6. Length of flagellomerus 2/length of flagellomerus 14: 2.0–2.2. Tarsal claws: simple. Metafemur length/width: 3.0–3.1. Metatibia inner spur length/metabasitarsus length: 0.4–0.5. Anteromesoscutum: mostly with deep, dense punctures (separated by less than 2.0 × its maximum diameter). Mesoscutellar disc: with punctures near margins, central part mostly smooth. Number of pits in scutoscutellar sulcus: 11 or 12. Maximum height of mesoscutellum lunules/maximum height of lateral face of mesoscutellum: 0.6–0.7. Propodeum areola: completely defined by carinae, including transverse carina extending to spiracle. Propodeum background sculpture: partly sculptured, especially on anterior 0.5. Mediotergite 1 length/width at posterior margin: 1.7–1.9. Mediotergite 1 shape: more or less parallel–sided. Mediotergite 1 sculpture: mostly sculptured, excavated area centrally with transverse striation inside and/or a polished knob centrally on posterior margin of mediotergite. Mediotergite 2 width at posterior margin/length: 4.0–4.3. Mediotergite 2 sculpture: mostly smooth. Outer margin of hypopygium: with a wide, medially folded, transparent, semi–desclerotized area; usually with 4 or more pleats. Ovipositor thickness: about same width throughout its length. Ovipositor sheaths length/metatibial length: 1.0–1.1. Length of fore wing veins r/2RS: 1.7–1.9. Length of fore wing veins 2RS/2M: 1.7–1.8. Length of fore wing veins 2M/(RS+M)b: 0.5–0.6. Pterostigma length/width: 2.6–3.0. Point of insertion of vein r in pterostigma: about half way point length of pterostigma. Angle of vein r with fore wing anterior margin: clearly outwards, inclined towards fore wing apex. Shape of junction of veins r and 2RS in fore wing: strongly angulated, sometimes with a knob.

Male. The only available specimen is in poor condition, missing metasoma, some legs and part of antennae.

Molecular data.

Sequences in BOLD: 3, barcode compliant sequences: 2.

Biology/ecology.

Solitary (Fig. 226). Hosts: Elachistidae, Antaeotricha similisEPR01, Stenoma Janzen07, Stenoma Janzen44; Crambidae, Omiodes Janzen03, Omiodes Janzen06.

Distribution.

Costa Rica, ACG.

Etymology.

We dedicate this species to Adrian Guadamuz in recognition of his diligent efforts for the ACG Programa de Parataxónomos and the plant inventory of ACG.

Apanteles aichagirardae Fernández-Triana, sp. n.

http://zoobank.org/4131A739-EEDF-41A0-8DC4-B52F162FAC83

http://species-id.net/wiki/Apanteles_aichagirardae

Fig. 35
Apanteles Rodriguez150 (Smith et al. 2006). Interim name provided by the authors.
Type locality.

COSTA RICA, Guanacaste, ACG, Sector Cacao, Sendero Derrumbe, 1220 meters, 10.92918, -85.46426.

Holotype.

♀ in CNC. Specimen labels: 1. COSTA RICA, Guanacaste, ACG, Sector Cacao, Sendero Derrumbe, 1220 meters, 24.iv.2006, 10.92918, -85.46426, 01-SRNP6737. 2. DHJPAR0012466.

Paratypes.

1 ♀ (CNC). COSTA RICA: Guanacaste, ACG database code: DHJPAR0012468.

Description.

Female. Body color: body mostly dark except for some sternites which may be pale. Antenna color: scape, pedicel, and flagellum dark. Coxae color (pro-, meso-, metacoxa): dark, dark, dark. Femora color (pro-, meso-, metafemur): pale, pale, dark. Tibiae color (pro-, meso-, metatibia): pale, pale, anteriorly pale/posteriorly dark. Tegula and humeral complex color: tegula pale, humeral complex half pale/half dark. Pterostigma color: dark. Fore wing veins color: mostly dark (a few veins may be unpigmented). Antenna length/body length: antenna about as long as body (head to apex of metasoma); if slightly shorter, at least extending beyond anterior 0.7 metasoma length. Body in lateral view: not distinctly flattened dorso–ventrally. Body length (head to apex of metasoma): 3.1–3.2 mm or 3.3–3.4 mm. Fore wing length: 3.3–3.4 mm or 3.5–3.6 mm. Ocular–ocellar line/posterior ocellus diameter: 1.7–1.9. Interocellar distance/posterior ocellus diameter: 1.7–1.9. Antennal flagellomerus 2 length/width: 3.2 or more. Antennal flagellomerus 14 length/width: 1.4–1.6. Length of flagellomerus 2/length of flagellomerus 14: 2.3–2.5. Tarsal claws: with single basal spine–like seta. Metafemur length/width: 3.0–3.1. Metatibia inner spur length/metabasitarsus length: 0.6–0.7. Anteromesoscutum: mostly with deep, dense punctures (separated by less than 2.0 × its maximum diameter). Mesoscutellar disc: mostly smooth. Number of pits in scutoscutellar sulcus: 9 or 10. Maximum height of mesoscutellum lunules/maximum height of lateral face of mesoscutellum: 0.4–0.5. Propodeum areola: completely defined by carinae, including transverse carina extending to spiracle. Propodeum background sculpture: mostly sculptured. Mediotergite 1 length/width at posterior margin: 1.1–1.3. Mediotergite 1 shape: clearly widening towards posterior margin. Mediotergite 1 sculpture: mostly sculptured, excavated area centrally with transverse striation inside and/or a polished knob centrally on posterior margin of mediotergite. Mediotergite 2 width at posterior margin/length: 3.2–3.5. Mediotergite 2 sculpture: with some sculpture, mostly near posterior margin. Outer margin of hypopygium: with a wide, medially folded, transparent, semi–desclerotized area; usually with 4 or more pleats. Ovipositor thickness: about same width throughout its length. Ovipositor sheaths length/metatibial length: 1.0–1.1. Length of fore wing veins r/2RS: 1.7–1.9. Length of fore wing veins 2RS/2M: 1.4–1.6. Length of fore wing veins 2M/(RS+M)b: 0.7–0.8. Pterostigma length/width: 2.6–3.0. Point of insertion of vein r in pterostigma: clearly beyond half way point length of pterostigma. Angle of vein r with fore wing anterior margin: more or less perpendicular to fore wing margin. Shape of junction of veins r and 2RS in fore wing: distinctly but not strongly angled.

Male. Unknown.

Molecular data.

Sequences in BOLD: 2, barcode compliant sequences: 2.

Biology/ecology.

Solitary. Host: Elachistidae, specimen with ACG database code: 01-SRNP-6737.

Distribution.

Costa Rica, ACG.

Comments.

This species is characterized by the combination of tegula different color from humeral complex, pterostigma brown, mediotergite 1 clearly widening towards posterior margin (1.3 × as long as wide at posterior margin), mediotergite 2 with posterior margin sinuate (width at expanded central area 1.7 × as large as width at lateral area), and ovipositor relatively thick (basal width about twice apical width).

Etymology.

We dedicate this species to Aicha Girardi, daughter of Caroline Boudreault (CNC, Ottawa) as an appreciation for Caroline’s support, especially photographing many types for this paper.

Type locality.

COSTA RICA, Alajuela, ACG, Sector Pitilla, Bullas, 440 meters, 10.98670, -85.38503.

Holotype.

♀ in CNC. Specimen labels: 1. DHJPAR0042048. 2. Voucher: D.H.Janzen & W.Hallwachs, DB: http://janzen.sas.upenn.edu, Area de Conservación Guanacaste, COSTA RICA, 11-SRNP-7011.

Paratypes.

6 ♀, 1 ♂ (CNC, NMNH). COSTA RICA: ACG database codes: DHJPAR0038184, DHJPAR0038224, DHJPAR0042041, DHJPAR0042043, DHJPAR0042044, DHJPAR0042062, DHJPAR0042425

Description.

Female. Body color: body mostly dark except for some sternites which may be pale. Antenna color: scape, pedicel, and flagellum dark. Coxae color (pro-, meso-, metacoxa): dark, dark, dark. Femora color (pro-, meso-, metafemur): pale, dark, dark. Tibiae color (pro-, meso-, metatibia): pale, pale, mostly dark but anterior 0.2 or less pale. Tegula and humeral complex color: tegula pale, humeral complex half pale/half dark. Pterostigma color: strongly white. Fore wing veins color: mostly white or entirely transparent. Antenna length/body length: antenna very short, barely or not extending beyond mesosoma length. Body in lateral view: not distinctly flattened dorso–ventrally. Body length (head to apex of metasoma): 2.3–2.4 mm. Fore wing length: 2.5–2.6 mm. Ocular–ocellar line/posterior ocellus diameter: 2.6 or more. Interocellar distance/posterior ocellus diameter: 2.0–2.2. Antennal flagellomerus 2 length/width: 1.7–1.9. Antennal flagellomerus 14 length/width: 1.1–1.3. Length of flagellomerus 2/length of flagellomerus 14: 1.7–1.9. Tarsal claws: simple. Metafemur length/width: 2.8–2.9. Metatibia inner spur length/metabasitarsus length: 0.4–0.5. Anteromesoscutum: mostly with shallow, dense punctures (separated by less than 2.0 × its maximum diameter). Mesoscutellar disc: mostly smooth. Number of pits in scutoscutellar sulcus: 7 or 8. Maximum height of mesoscutellum lunules/maximum height of lateral face of mesoscutellum: 0.6–0.7. Propodeum areola: completely defined by carinae, including transverse carina extending to spiracle. Propodeum background sculpture: partly sculptured, especially on anterior 0.5. Mediotergite 1 length/width at posterior margin: 2.9–3.1. Mediotergite 1 shape: mostly parallel–sided for 0.5–0.7 of its length, then narrowing posteriorly so mediotergite anterior width >1.1 × posterior width. Mediotergite 1 sculpture: mostly smooth. Mediotergite 2 width at posterior margin/length: 3.2–3.5. Mediotergite 2 sculpture: mostly smooth. Outer margin of hypopygium: with a wide, medially folded, transparent, semi–desclerotized area; usually with 4 or more pleats. Ovipositor thickness: anterior width 3.0–5.0 × posterior width (beyond ovipositor constriction). Ovipositor sheaths length/metatibial length: 0.4–0.5. Length of fore wing veins r/2RS: 1.7–1.9. Length of fore wing veins 2RS/2M: 1.1–1.3. Length of fore wing veins 2M/(RS+M)b: 0.5–0.6. Pterostigma length/width: 2.6–3.0. Point of insertion of vein r in pterostigma: about half way point length of pterostigma. Angle of vein r with fore wing anterior margin: more or less perpendicular to fore wing margin. Shape of junction of veins r and 2RS in fore wing: distinctly but not strongly angled.

Male. The specimen available for study is in poor condition, but resemble the females.

Molecular data.

Sequences in BOLD: 10, barcode compliant sequences: 10.

Biology/ecology.

Solitary. Hosts: Crambidae, Omiodes cuniculalis, Prenesta Janzen196.

Comments.

This species is characterized by a very distinctive hypopygium (with a relatively wide fold where no pleats are visible), ovipositor sheaths (very short and shaped as a broad spatula) and ovipositor (short and strongly curved downwards); it is further distinguished by antenna much shorter than body, white pterostigma, white or transparent fore wing veins, and elongate glossa. The unique hypopygium, ovipositor sheaths, and ovipositor, suggest that this species may be placed in a new genus when there are more studies on the phylogeny of Microgastrinae. Because that is beyond the scope of this paper, we describe this species in Apanteles.

Etymology.

We dedicate this species to Aida López in recognition of her diligent efforts in the Programa del Comedor Santa Rosa.

Type locality.

COSTA RICA, Guanacaste, ACG, Sector Cacao, Sendero Cima, 1460m, 10.93328, -85.45729.

Holotype.

♀ in CNC. Specimen labels: 1. DHJPAR0012506. 2. 24–31 Aug. 1998, CLC.

Description.

Body color: head and mesosoma mostly dark, metasoma with some tergites and/or most of sternites pale. Antenna color: scape, pedicel, and flagellum dark. Coxae color (pro-, meso-, metacoxa): pale, pale, dark. Femora color (pro-, meso-, metafemur): pale, pale, mostly pale but with dark area dorsally. Tibiae color (pro-, meso-, metatibia): pale, pale, anteriorly pale/posteriorly dark. Tegula and humeral complex color: both pale. Pterostigma color: dark. Fore wing veins color: mostly dark (a few veins may be unpigmented). Antenna length/body length: antenna about as long as body (head to apex of metasoma); if slightly shorter, at least extending beyond anterior 0.7 metasoma length. Body in lateral view: not distinctly flattened dorso–ventrally. Body length (head to apex of metasoma): 2.7–2.8 mm. Fore wing length: 2.9–3.0 mm. Ocular–ocellar line/posterior ocellus diameter: 2.3–2.5. Interocellar distance/posterior ocellus diameter: 2.0–2.2. Antennal flagellomerus 2 length/width: 2.3–2.5. Antennal flagellomerus 14 length/width: 1.4–1.6. Length of flagellomerus 2/length of flagellomerus 14: 2.0–2.2. Tarsal claws: simple. Metafemur length/width: 3.2–3.3. Metatibia inner spur length/metabasitarsus length: 0.4–0.5. Anteromesoscutum: mostly with deep, dense punctures (separated by less than 2.0 × its maximum diameter). Mesoscutellar disc: mostly punctured. Number of pits in scutoscutellar sulcus: 5 or 6. Maximum height of mesoscutellum lunules/maximum height of lateral face of mesoscutellum: 0.4–0.5. Propodeum areola: completely defined by carinae, including transverse carina extending to spiracle. Propodeum background sculpture: mostly sculptured. Mediotergite 1 length/width at posterior margin: 2.3–2.5. Mediotergite 1 shape: mostly parallel–sided for 0.5–0.7 of its length, then narrowing posteriorly so mediotergite anterior width >1.1 × posterior width. Mediotergite 1 sculpture: mostly sculptured, excavated area centrally with transverse striation inside and/or a polished knob centrally on posterior margin of mediotergite. Mediotergite 2 width at posterior margin/length: 2.8–3.1. Mediotergite 2 sculpture: more or less fully sculptured, with longitudinal striation. Outer margin of hypopygium: with a wide, medially folded, transparent, semi–desclerotized area; usually with 4 or more pleats. Ovipositor thickness: about same width throughout its length. Ovipositor sheaths length/metatibial length: 1.0–1.1. Length of fore wing veins r/2RS: 1.4–1.6. Length of fore wing veins 2RS/2M: 0.8 or less. Length of fore wing veins 2M/(RS+M)b: 1.1–1.3. Pterostigma length/width: 2.6–3.0. Point of insertion of vein r in pterostigma: about half way point length of pterostigma. Angle of vein r with fore wing anterior margin: more or less perpendicular to fore wing margin. Shape of junction of veins r and 2RS in fore wing: distinctly but not strongly angled.

Male. Unknown.

Molecular data.

Sequences in BOLD: 4, barcode compliant sequences: 4.

Biology/ecology.

Host: Malaise-trapped.

Distribution.

Costa Rica, ACG.

Etymology.

We dedicate this species to Alban Jiménez in recognition of his diligent efforts for the ACG Programa de Educacion Biológica.

Apanteles albinervis (Cameron, 1904), stat. rev. Fig. 164
Urogaster albinervis Cameron, 1904: 261.
Apanteles albinervican Shenefelt, 1972: 438. Invalid replacement name.
Type locality.

MEXICO.

Holotype.

♂, BMNH (examined).

Description.

Male. Body color: body mostly dark except for some sternites which may be pale. Antenna color: scape, pedicel, and flagellum dark. Coxae color (pro-, meso-, metacoxa): dark, dark, dark (?). Femora color (pro-, meso-, metafemur): anteriorly dark/posteriorly pale, dark, dark. Tibiae color (pro-, meso-, metatibia): pale, anteriorly pale/posteriorly dark, anteriorly pale/posteriorly dark. Tegula and humeral complex color: tegula pale, humeral complex half pale/half dark. Pterostigma color: mostly pale and/or transparent, with thin dark borders. Fore wing veins color: mostly white or entirely transparent. Antenna length/body length: antenna about as long as body (head to apex of metasoma); if slightly shorter, at least extending beyond anterior 0.7 metasoma length (?). Body in lateral view: not distinctly flattened dorso–ventrally. Body length (head to apex of metasoma): 2.3–2.4 mm. Fore wing length: 2.5–2.6 mm. Ocular–ocellar line/posterior ocellus diameter: 2.0–2.2. Interocellar distance/posterior ocellus diameter: 1.7–1.9. Tarsal claws: simple (?). Metafemur length/width: 3.0–3.1. Metatibia inner spur length/metabasitarsus length: 0.4–0.5. Anteromesoscutum: mostly with shallow, dense punctures (separated by less than 2.0 × its maximum diameter). Mesoscutellar disc: mostly smooth. Number of pits in scutoscutellar sulcus: 9 or 10. Maximum height of mesoscutellum lunules/maximum height of lateral face of mesoscutellum: 0.6–0.7. Propodeum areola: completely defined by carinae, including transverse carina extending to spiracle. Propodeum background sculpture: partly sculptured, especially on anterior 0.5. Mediotergite 1 length/width at posterior margin: 1.4–1.6. Mediotergite 1 shape: slightly widening from anterior margin to 0.7–0.8 mediotergite length (where maximum width is reached), then narrowing towards posterior margin. Mediotergite 1 sculpture: with some sculpture near lateral margins and/or posterior 0.2–0.4 of mediotergite. Mediotergite 2 width at posterior margin/length: 2.8–3.1. Mediotergite 2 sculpture: mostly smooth. Length of fore wing veins r/2RS: 1.7–1.9. Length of fore wing veins 2RS/2M: 1.4–1.6. Length of fore wing veins 2M/(RS+M)b: 0.5–0.6. Pterostigma length/width: 3.6 or more. Point of insertion of vein r in pterostigma: about half way point length of pterostigma. Angle of vein r with fore wing anterior margin: more or less perpendicular to fore wing margin. Shape of junction of veins r and 2RS in fore wing: distinctly but not strongly angled.

Female. Unknown.

Molecular data.

No molecular data available for this species.

Biology/ecology.

Nothing is known of its hosts.

Distribution.

Known only from the male holotype, which was collected in “Mexico”. There is no suggestion that this species occurs in Costa Rica or ACG.

Comments.

The history of the name “Apanteles albinervis” needs clarification. Cameron (1904) described the species “Urogaster albinervis” from Mexico. Urogaster was later synonymized under Apanteles by Szépligeti (1904), but Cameron’s species was not formally transferred to it until Shenefelt (1972)’s World Catalogue of Hymenoptera. In the meantime, Tobias (1964) had described a species from Kazakhstan as “Apanteles albinervis” – later found to be widely distributed in the Palearctic region (Yu et al. 2005) and not related at all to the Mexican species. Acting as the first reviser, Shenefelt realized the problem of a secondary homonym, but mistakenly assigned a replacement name for the oldest (Cameron 1904) instead of the youngest (Tobias 1964) name. As a result, Apanteles albinervican Shenefelt, 1972 became a replacement name for Urogaster albinervis Cameron, 1904, while Apanteles albinervis Tobias, 1964 remained unchanged (Shenefelt 1972). Article 24.2.5 of the International Code of Zoological Nomenclature (ICZN 1999) regulates “Unnecessary action by a First Reviser” and states that “if it is shown subsequently that the precedence of names, spellings or acts can be objectively determined, the action of the First Reviser is nullified”. Thus we consider here Apanteles albinervican Shenefelt, 1972 an invalid replacement name for Apanteles albinervis (Cameron, 1904) and reinstate the latter name. For details of the revised status of the Tobias species see section “Species excluded from Apanteles”.

Another unrelated use of the name “Apanteles albinervis”, was by Ashmead (1905), who described a species from the Philippines as “Urogaster albinervis”. That became a primary homonym of Urogaster albinervis Cameron; however, a replacement name for the Philipine species, Apanteles lucidinervis, was provided by Wilkinson (1928).

Type locality.

COSTA RICA, Guanacaste, ACG, Sector El Hacha, Sendero Bejuquilla, 280 m, 11.03004, -85.52699.

Holotype.

♀ in CNC. Specimen labels: 1. DHJPAR0012499. 2. COSTA RICA, Guanacaste, ACG, Sector El Hacha, Sendero Bejuquilla, 280 meters, 11.03004 Longitude: -85.52699, D.H. Janzen &♀ in CNC. Specimen labels: 1. DHJPAR0012499. 2. COSTA RICA, Guanacaste, ACG, Sector El Hacha, Sendero Bejuquilla, 280 meters, 11.03004 Longitude: -85.52699, D.H. Janzen & W. Hallwachs..

Description.

Female. Body color: head pale, mesosoma extensively pale (anteromesoscutum and scutellar disc). Antenna color: scape and/or pedicel pale, flagellum dark. Coxae color (pro-, meso-, metacoxa): pale, pale, partially pale/partially dark. Femora color (pro-, meso-, metafemur): pale, pale, mostly pale but posterior 0.2 or less dark. Tibiae color (pro-, meso-, metatibia): pale, pale, mostly pale but with posterior 0.2 or less dark. Tegula and humeral complex color: both pale. Pterostigma color: dark. Fore wing veins color: mostly dark (a few veins may be unpigmented). Body in lateral view: not distinctly flattened dorso–ventrally. Body length (head to apex of metasoma): 3.1–3.2 mm. Fore wing length: 3.3–3.4 mm. Ocular–ocellar line/posterior ocellus diameter: 2.3–2.5. Interocellar distance/posterior ocellus diameter: 2.3–2.5. Antennal flagellomerus 2 length/width: 2.6–2.8. Antennal flagellomerus 14 length/width: 1.7–1.9. Length of flagellomerus 2/length of flagellomerus 14: 2.3–2.5. Tarsal claws: with single basal spine–like seta. Metafemur length/width: 2.8–2.9. Anteromesoscutum: mostly with deep, dense punctures (separated by less than 2.0 × its maximum diameter). Mesoscutellar disc: mostly punctured. Number of pits in scutoscutellar sulcus: 5 or 6. Maximum height of mesoscutellum lunules/maximum height of lateral face of mesoscutellum: 0.4–0.5. Propodeum areola: partially defined by carinae on posterior 0.3–0.5 of its length, widely open anteriorly. Propodeum background sculpture: mostly sculptured. Mediotergite 1 length/width at posterior margin: 2.6–2.8. Mediotergite 1 shape: mostly parallel–sided for 0.5–0.7 of its length, then narrowing posteriorly so mediotergite anterior width >1.1 × posterior width. Mediotergite 1 sculpture: mostly sculptured, excavated area centrally with transverse striation inside and/or a polished knob centrally on posterior margin of mediotergite. Mediotergite 2 width at posterior margin/length: 3.2–3.5. Mediotergite 2 sculpture: with some sculpture, mostly near posterior margin. Outer margin of hypopygium: with a wide, medially folded, transparent, semi–desclerotized area; usually with 4 or more pleats. Ovipositor thickness: anterior width at most 2.0 × posterior width (beyond ovipositor constriction). Ovipositor sheaths length/metatibial length: 0.8–0.9. Length of fore wing veins r/2RS: 2.3 or more. Length of fore wing veins 2RS/2M: 0.9–1.0. Length of fore wing veins 2M/(RS+M)b: 0.7–0.8. Pterostigma length/width: 2.6–3.0. Point of insertion of vein r in pterostigma: about half way point length of pterostigma. Angle of vein r with fore wing anterior margin: clearly outwards, inclined towards fore wing apex. Shape of junction of veins r and 2RS in fore wing: distinctly but not strongly angled.

Male. Unknown.

Molecular data.

Sequences in BOLD: 1, barcode compliant sequences: 1.

Biology/ecology.

Malaise trapped.

Distribution.

Costa Rica, ACG.

Comments.

This species is very distinctive, characterized by head and most of mediotergite 1 orange, mediotergite 3 partially yellow, and mesoscutellar disc mostly punctured.

Etymology.

We dedicate this species to Alejandro Masis in recognition of his diligent efforts to administrate and protect the entire ACG.

Type locality.

COSTA RICA, Alajuela, ACG, Sector Rincon Rain Forest, Sendero Albergue Crater, 980m, 10.84886, -85.3281.

Holotype.

♀ in CNC. Specimen labels: 1. Voucher: D.H.Janzen & W.Hallwachs, DB: http://janzen.sas.upenn.edu, Area de Conservación Guanacaste, COSTA RICA, 09-SRNP-4936. 2. DHJPAR0039759.

Paratypes.

3 ♀, 3 ♂ (BMNH, CNC, INBIO, INHS, NMNH). COSTA RICA, ACG database codes: DHJPAR0027612, DHJPAR0035523, DHJPAR0038321, DHJPAR0039030, DHJPAR0039734, DHJPAR0039775.

Description.

Female. Body color: body mostly dark except for some sternites which may be pale. Antenna color: scape, pedicel, and flagellum dark. Coxae color (pro-, meso-, metacoxa): dark, dark, dark. Femora color (pro-, meso-, metafemur): anteriorly dark/posteriorly pale, dark, dark. Tibiae color (pro-, meso-, metatibia): pale, pale, anteriorly pale/posteriorly dark. Tegula and humeral complex color: tegula pale, humeral complex half pale/half dark. Pterostigma color: dark. Fore wing veins color: mostly dark (a few veins may be unpigmented). Antenna length/body length: antenna about as long as body (head to apex of metasoma); if slightly shorter, at least extending beyond anterior 0.7 metasoma length. Body in lateral view: not distinctly flattened dorso–ventrally. Body length (head to apex of metasoma): 3.3–3.4 mm or 3.5–3.6 mm. Fore wing length: 3.3–3.4 mm, 3.5–3.6 mm or 3.7–3.8 mm. Ocular–ocellar line/posterior ocellus diameter: 2.3–2.5. Interocellar distance/posterior ocellus diameter: 1.4–1.6. Antennal flagellomerus 2 length/width: 2.9–3.1. Antennal flagellomerus 14 length/width: 1.7–1.9. Length of flagellomerus 2/length of flagellomerus 14: 2.3–2.5. Tarsal claws: with single basal spine–like seta. Metafemur length/width: 3.0–3.1. Metatibia inner spur length/metabasitarsus length: 0.4–0.5. Anteromesoscutum: mostly with deep, dense punctures (separated by less than 2.0 × its maximum diameter). Mesoscutellar disc: mostly smooth. Number of pits in scutoscutellar sulcus: 11 or 12. Maximum height of mesoscutellum lunules/maximum height of lateral face of mesoscutellum: 0.6–0.7. Propodeum areola: completely defined by carinae, including transverse carina extending to spiracle. Propodeum background sculpture: mostly sculptured. Mediotergite 1 length/width at posterior margin: 2.3–2.5. Mediotergite 1 shape: mostly parallel–sided for 0.5–0.7 of its length, then narrowing posteriorly so mediotergite anterior width >1.1 × posterior width. Mediotergite 1 sculpture: mostly sculptured, excavated area centrally with transverse striation inside and/or a polished knob centrally on posterior margin of mediotergite. Mediotergite 2 width at posterior margin/length: 2.0–2.3. Mediotergite 2 sculpture: mostly smooth. Outer margin of hypopygium: with a wide, medially folded, transparent, semi–desclerotized area; usually with 4 or more pleats. Ovipositor thickness: about same width throughout its length. Ovipositor sheaths length/metatibial length: 1.8–1.9. Length of fore wing veins r/2RS: 1.7–1.9. Length of fore wing veins 2RS/2M: 1.9–2.0. Length of fore wing veins 2M/(RS+M)b: 0.5–0.6. Pterostigma length/width: 3.6 or more. Point of insertion of vein r in pterostigma: clearly beyond half way point length of pterostigma. Angle of vein r with fore wing anterior margin: more or less perpendicular to fore wing margin. Shape of junction of veins r and 2RS in fore wing: distinctly but not strongly angled.

Male. Similar to female, except for mediotergite 2 much less quadrate (i.e., much more transverse).

Molecular data.

Sequences in BOLD: 11, barcode compliant sequences: 10.

Biology/ecology.

Solitary (Fig. 234). Hosts: Elachistidae, Antaeotricha Janzen106, Antaeotricha Janzen366, Lethata trochalosticta.

Distribution.

Costa Rica, ACG.

Etymology.

We dedicate this species to Alejandro Mora in recognition of his diligent efforts for the ACG administration and Programa de Contabilidad.

Apanteles alvarougaldei Fernández-Triana, sp. n.

http://zoobank.org/A19B70E1-1448-4C66-82EA-5C3E7F0A94C7

http://species-id.net/wiki/Apanteles_alvarougaldei

Figs 167, 304
Apanteles Rodriguez40 (Smith et al. 2006). Interim name provided by the authors.
Type locality.

COSTA RICA, Alajuela, ACG, Sector San Cristobal, Puente Palma, 460m, 10.9163, -85.37869.

Holotype.

♀ in CNC. Specimen labels: 1. COSTA RICA, Alajuela, ACG, Sector San Cristobal, Puente Palma, 460m, 23.iv.2006, 10.9163, -85.37869, DHJPAR0011962.

Paratypes.

22 ♀, 9 ♂ (BMNH, CNC, INBIO, INHS, NMNH). COSTA RICA, ACG database codes: DHJPAR0001584, DHJPAR0002680, DHJPAR0003037, DHJPAR0011962, 93-SRNP-4213.

Description.

Female. Metatibia color (outer face): entirely or mostly (>0.7 metatibia length) dark brown to black, with yellow to white coloration usually restricted to anterior 0.2 or less. Fore wing veins color: veins C+Sc+R and R1 with brown coloration restricted narrowly to borders, interior area of those veins and pterostigma (and sometimes veins r, 2RS and 2M) transparent or white; other veins mostly transparent. Antenna length/body length: antenna about as long as body (head to apex of metasoma); if slightly shorter, at least extending beyond anterior 0.7 metasoma length. Body length (head to apex of metasoma): 2.3–2.4 mm, 2.5–2.6 mm, rarely 2.1–2.2 mm. Fore wing length: 2.5–2.6 mm or 2.7–2.8 mm. Metafemur length/width: 2.6–2.7, 2.8–2.9 or 3.0–3.1. Mediotergite 1 length/width at posterior margin: 2.3–2.4. Mediotergite 1 maximum width/width at posterior margin: 1.6–1.7. Ovipositor sheaths length/metafemur length: 0.9 or 1.0. Ovipositor sheaths length/metatibia length: 0.8 or 0.9.

Molecular data.

Sequences in BOLD: 8, barcode compliant sequences: 7.

Biology/ecology.

Gregarious (Fig. 304). Host: Hesperiidae, Bungalotis quadratum.

Distribution.

Costa Rica, ACG.

Etymology.

We dedicate this species to Alvaro Ugalde in recogniton of his diligent efforts in founding and guiding the National Park System of Costa Rica.

Apanteles anabellecordobae Fernández-Triana, sp. n.

http://zoobank.org/80ACA979-8A0B-4EA2-A5C1-D33A871686F5

http://species-id.net/wiki/Apanteles_anabellecordobae

Figs 51, 244
Apanteles Rodriguez05 (Smith et al. 2006). Interim name provided by the authors.
Type locality.

COSTA RICA, Guanacaste, ACG, Sector Cacao, Sendero Circular, 1185m, 10.92714, -85.46683.

Holotype.

♀ in CNC. Specimen labels: 1. COSTA RICA, Guanacaste, ACG, Sector Cacao, Sendero Circular, 01/05/2001, Mariano Pereira. 2. 01-SRNP-6021, Achlyodes selva, Zanthoxylum melanostictum. 3. DHJPAR0001550.

Paratypes.

68 ♀, 11 ♂ (BMNH, CNC, INBIO, INHS, NMNH). COSTA RICA: ACG database codes: See Appendix 2 for detailed label data.

Description.

Female. Body color: body mostly dark except for some sternites which may be pale. Antenna color: scape, pedicel, and flagellum dark. Coxae color (pro-, meso-, metacoxa): dark, dark, dark. Femora color (pro-, meso-, metafemur): anteriorly dark/posteriorly pale, dark, dark. Tibiae color (pro-, meso-, metatibia): pale, pale, anteriorly pale/posteriorly dark. Tegula and humeral complex color: tegula pale, humeral complex dark. Pterostigma color: dark with pale spot at base. Fore wing veins color: mostly dark (a few veins may be unpigmented). Antenna length/body length: antenna about as long as body (head to apex of metasoma); if slightly shorter, at least extending beyond anterior 0.7 metasoma length. Body in lateral view: not distinctly flattened dorso–ventrally. Body length (head to apex of metasoma): 2.7–2.8 mm, 2.9–3.0 mm or 3.1–3.2 mm. Fore wing length: 2.9–3.0 mm, 3.1–3.2 mm or 3.3–3.4 mm. Ocular–ocellar line/posterior ocellus diameter: 1.7–1.9. Interocellar distance/posterior ocellus diameter: 1.4–1.6. Antennal flagellomerus 2 length/width: 2.6–2.8. Antennal flagellomerus 14 length/width: 1.4–1.6. Length of flagellomerus 2/length of flagellomerus 14: 2.0–2.2. Tarsal claws: with single basal spine–like seta. Metafemur length/width: 3.0–3.1. Metatibia inner spur length/metabasitarsus length: 0.4–0.5. Anteromesoscutum: mostly with deep, dense punctures (separated by less than 2.0 × its maximum diameter). Mesoscutellar disc: with punctures near margins, central part mostly smooth. Number of pits in scutoscutellar sulcus: 9 or 10. Maximum height of mesoscutellum lunules/maximum height of lateral face of mesoscutellum: 0.6–0.7. Propodeum areola: completely defined by carinae, including transverse carina extending to spiracle. Propodeum background sculpture: mostly sculptured. Mediotergite 1 length/width at posterior margin: 4.1 or more. Mediotergite 1 shape: mostly parallel–sided for 0.5–0.7 of its length, then narrowing posteriorly so mediotergite anterior width >1.1 × posterior width. Mediotergite 1 sculpture: with some sculpture near lateral margins and/or posterior 0.2–0.4 of mediotergite. Mediotergite 2 width at posterior margin/length: 2.8–3.1. Mediotergite 2 sculpture: mostly smooth. Outer margin of hypopygium: inflexible (without a folded, transparent, semi–desclerotized area); with no pleats visible. Ovipositor thickness: anterior width 3.0–5.0 × posterior width (beyond ovipositor constriction). Ovipositor sheaths length/metatibial length: 0.8–0.9. Length of fore wing veins r/2RS: 2.3 or more. Length of fore wing veins 2RS/2M: 1.1–1.3. Length of fore wing veins 2M/(RS+M)b: 0.7–0.8. Pterostigma length/width: 3.1–3.5. Point of insertion of vein r in pterostigma: about half way point length of pterostigma. Angle of vein r with fore wing anterior margin: more or less perpendicular to fore wing margin. Shape of junction of veins r and 2RS in fore wing: distinctly but not strongly angled.

Male. Similar to female.

Molecular data.

Sequences in BOLD: 29, barcode compliant sequences: 24, haplotypes: 2.

Biology/ecology.

Gregarious (Fig. 244). Hosts: Hesperiidae, Achlyodes busirus, Achlyodes pallida.

Distribution.

Costa Rica, ACG.

Coments.

Five specimens (4 ♀ and 1 ♂ )were reared from the same Achlyodes pallida caterpillar as the barcoded specimen DHJPAR0005308 (all of them with ACG code 97-SRNP-984). The sequenced specimen clusters apart from the rest of the species, differing by 2.45 % base pairs. Additionally, the locality for specimens with code 97-SRNP-984 is at a lower altitude (90m) compared with the rest of the species (mostly found between 500-1140 m, with only four specimens between 280-290m). Those five specimens might represent a different species, but lacking the support of morphological data and host records (which do not seem to differ) we have decided to keep them under this species for the time being. We have, however, excluded them from the paratypes series.

Etymology.

We dedicate this species to Anabelle Cordoba in recognition of her diligent efforts for the ACG Programa de Parataxónomos and Estación Biológica Caribe of ACG.

Type locality.

COSTA RICA, Guanacaste, ACG,

Holotype.

♀ in CNC. Specimen labels: 1. DHJPAR0041897.

Paratypes.

2 ♀ (CNC). COSTA RICA: Guanacaste, ACG database code: DHJPAR0041983.

Description.

Female. Body color: body mostly dark except for some sternites which may be pale. Antenna color: scape, pedicel, and flagellum dark. Coxae color (pro-, meso-, metacoxa): dark, dark, dark. Femora color (pro-, meso-, metafemur): pale, pale, mostly pale but posterior 0.2 or less dark. Tibiae color (pro-, meso-, metatibia): pale, pale, mostly pale but with posterior 0.2 or less dark. Tegula and humeral complex color: both pale. Pterostigma color: dark with pale spot at base. Fore wing veins color: partially pigmented (a few veins may be dark but most are pale). Antenna length/body length: antenna about as long as body (head to apex of metasoma); if slightly shorter, at least extending beyond anterior 0.7 metasoma length. Body in lateral view: not distinctly flattened dorso–ventrally. Body length (head to apex of metasoma): 3.3–3.4 mm. Fore wing length: 3.5–3.6 mm. Ocular–ocellar line/posterior ocellus diameter: 2.0–2.2. Interocellar distance/posterior ocellus diameter: 1.7–1.9. Antennal flagellomerus 2 length/width: 2.9–3.1. Antennal flagellomerus 14 length/width: 1.7–1.9. Length of flagellomerus 2/length of flagellomerus 14: 2.3–2.5. Tarsal claws: simple. Metafemur length/width: 3.0–3.1. Metatibia inner spur length/metabasitarsus length: 0.4–0.5. Anteromesoscutum: mostly with deep, dense punctures (separated by less than 2.0 × its maximum diameter). Mesoscutellar disc: mostly smooth. Number of pits in scutoscutellar sulcus: 11 or 12. Maximum height of mesoscutellum lunules/maximum height of lateral face of mesoscutellum: 0.6–0.7. Propodeum areola: completely defined by carinae, including transverse carina extending to spiracle. Propodeum background sculpture: mostly sculptured. Mediotergite 1 length/width at posterior margin: 1.7–1.9. Mediotergite 1 shape: more or less parallel–sided. Mediotergite 1 sculpture: mostly sculptured, excavated area centrally with transverse striation inside and/or a polished knob centrally on posterior margin of mediotergite. Mediotergite 2 width at posterior margin/length: 2.8–3.1. Mediotergite 2 sculpture: mostly smooth. Outer margin of hypopygium: with a wide, medially folded, transparent, semi–desclerotized area; usually with 4 or more pleats. Ovipositor thickness: about same width throughout its length. Ovipositor sheaths length/metatibial length: 2.0 or more. Length of fore wing veins r/2RS: 2.0–2.2. Length of fore wing veins 2RS/2M: 1.4–1.6. Length of fore wing veins 2M/(RS+M)b: 0.5–0.6. Pterostigma length/width: 3.1–3.5. Point of insertion of vein r in pterostigma: clearly beyond half way point length of pterostigma. Angle of vein r with fore wing anterior margin: more or less perpendicular to fore wing margin. Shape of junction of veins r and 2RS in fore wing: distinctly but not strongly angled.

Male. Unknown.

Molecular data.

Sequences in BOLD: 6, barcode compliant sequences: 6.

Biology/ecology.

Solitary. Hosts: Elachistidae, elachjanzen01 Janzen120, Antaeotricha Janzen146, Oecophoridae, Inga biolep146DHJ01.

Distribution.

Costa Rica, ACG.

Etymology.

We dedicate this species to Ana Marenco in recognition of her diligent efforts for the ACG office administration in Sector Santa Rosa.

Apanteles anamartinezae Fernández-Triana, sp. n.

http://zoobank.org/9BE69016-4E47-4E77-8684-E583304C8932

http://species-id.net/wiki/Apanteles_anamartinezae

Figs 25, 227
Apanteles Rodriguez51 (Smith et al. 2006). Interim name provided by the authors.
Type locality.

COSTA RICA, Alajuela, ACG, Sector San Cristobal, Rio Blanco Abajo, 500m, 10.90037, -85.37254.

Holotype.

♀ in CNC. Specimen labels: 1. Costa Rica: Alajuela, ACG, Sector San Cristobal, Rio Blanco Abajo, 15.ii.2007, 500m, 10.90037, -85.37254, DHJPAR0012998.

Paratypes.

22 ♀, 9 ♂ (BMNH, CNC, INBIO, INHS, NMNH). COSTA RICA, ACG database codes: DHJPAR0002154, DHJPAR0002227, DHJPAR0004974 DHJPAR0012976, DHJPAR0020130, DHJPAR0031090, 99-SRNP-4439, 03-SRNP-5327.

Description.

Female. Body color: body mostly dark except for some sternites which may be pale. Antenna color: scape, pedicel, and flagellum dark. Coxae color (pro-, meso-, metacoxa): dark, dark, dark. Femora color (pro-, meso-, metafemur): anteriorly dark/posteriorly pale, dark, dark. Tibiae color (pro-, meso-, metatibia): pale, anteriorly pale/posteriorly dark, dark. Tegula and humeral complex color: tegula pale, humeral complex half pale/half dark. Pterostigma color: mostly pale and/or transparent, with thin dark borders. Fore wing veins color: partially pigmented (a few veins may be dark but most are pale). Antenna length/body length: antenna about as long as body (head to apex of metasoma); if slightly shorter, at least extending beyond anterior 0.7 metasoma length. Body in lateral view: not distinctly flattened dorso–ventrally. Body length (head to apex of metasoma): 2.1–2.2 mm, 2.3–2.4 mm or 2.5–2.6 mm. Fore wing length: 2.1–2.2 mm, 2.3–2.4 mm or 2.5–2.6 mm. Ocular–ocellar line/posterior ocellus diameter: 2.0–2.2. Interocellar distance/posterior ocellus diameter: 1.7–1.9. Antennal flagellomerus 2 length/width: 2.0–2.2. Antennal flagellomerus 14 length/width: 1.7–1.9. Length of flagellomerus 2/length of flagellomerus 14: 3.2 or more. Tarsal claws: with single basal spine–like seta. Metafemur length/width: 3.4–3.5. Metatibia inner spur length/metabasitarsus length: 0.4–0.5. Anteromesoscutum: mostly with deep, dense punctures (separated by less than 2.0 × its maximum diameter). Mesoscutellar disc: mostly smooth. Number of pits in scutoscutellar sulcus: 7 or 8. Maximum height of mesoscutellum lunules/maximum height of lateral face of mesoscutellum: 0.6–0.7. Propodeum areola: completely defined by carinae, including transverse carina extending to spiracle. Propodeum background sculpture: partly sculptured, especially on anterior 0.5. Mediotergite 1 length/width at posterior margin: 1.4–1.6. Mediotergite 1 shape: more or less parallel–sided. Mediotergite 1 sculpture: mostly sculptured, excavated area centrally with transverse striation inside and/or a polished knob centrally on posterior margin of mediotergite. Mediotergite 2 width at posterior margin/length: 4.0–4.3. Mediotergite 2 sculpture: mostly smooth. Outer margin of hypopygium: with a wide, medially folded, transparent, semi–desclerotized area; usually with 4 or more pleats. Ovipositor thickness: about same width throughout its length. Ovipositor sheaths length/metatibial length: 0.6–0.7 or 0.8–0.9. Length of fore wing veins r/2RS: 1.7–1.9. Length of fore wing veins 2RS/2M: 1.4–1.6. Length of fore wing veins 2M/(RS+M)b: 0.7–0.8. Pterostigma length/width: 3.1–3.5. Point of insertion of vein r in pterostigma: about half way point length of pterostigma. Angle of vein r with fore wing anterior margin: clearly outwards, inclined towards fore wing apex. Shape of junction of veins r and 2RS in fore wing: distinctly but not strongly angled.

Male. As in female, with slender mediotergite 1.

Molecular data.

Sequences in BOLD: 101, barcode compliant sequences: 98.

Biology/ecology.

Gregarious (Fig. 227). Hosts: Attevidae, Atteva aurea, Atteva pustulella, Atteva zebra.

Distribution.

Costa Rica, ACG.

Comments.

Although this species is clearly gregarious, in a few cases only one wasp cocoon is encountered, owing to the small size of the host caterpillar, which may support just one wasp larva, or just an artefact of the lightly silked cocoons falling apart, thus reducing the actual number of cocoons per caterpillar that are encountered when sampling.

Etymology.

We dedicate this species to Ana Martínez in recognition of her diligent efforts for the ACG Programa de Contabilidad.

Apanteles anapiedrae Fernández-Triana, sp. n.

http://zoobank.org/92E5C788-4C54-45AC-8092-E3A8EDBDFBA2

http://species-id.net/wiki/Apanteles_anapiedrae

Figs 67, 257
Apanteles Rodriguez156 (Smith et al. 2006). Interim name provided by the authors.
Type locality.

COSTA RICA, Alajuela, ACG, Sector San Cristobal, Finca San Gabriel, 645m, 10.87766, -85.39343.

Holotype.

♀ in CNC. Specimen labels: 1. COSTA RICA, ACG, Sector San Cristobal, Finca S. Gabriel, 645m, DHJPAR0039721.

Paratypes.

17 ♀, 5 ♂ (BMNH, CNC, INBIO, INHS, NMNH). COSTA RICA, Alajuela, ACG database codes: DHJPAR0039721, 09-SRNP-3890, 10-SRNP-1054.

Description.

Female. Body color: body mostly dark except for some sternites which may be pale. Antenna color: scape and/or pedicel pale, flagellum dark. Coxae color (pro-, meso-, metacoxa): dark, dark, dark. Femora color (pro-, meso-, metafemur): anteriorly dark/posteriorly pale, dark, dark. Tibiae color (pro-, meso-, metatibia): pale, pale, dark. Tegula and humeral complex color: both dark. Pterostigma color: dark. Fore wing veins color: mostly dark (a few veins may be unpigmented). Antenna length/body length: antenna shorter than body (head to apex of metasoma), not extending beyond anterior 0.7 metasoma length. Body in lateral view: distinctly flattened dorso–ventrally. Body length (head to apex of metasoma): 2.0 mm or less. Fore wing length: 2.0 mm or less. Ocular–ocellar line/posterior ocellus diameter: 2.6 or more. Interocellar distance/posterior ocellus diameter: 1.4–1.6. Antennal flagellomerus 2 length/width: 2.0–2.2. Antennal flagellomerus 14 length/width: 1.4–1.6. Length of flagellomerus 2/length of flagellomerus 14: 1.7–1.9. Tarsal claws: simple. Metafemur length/width: 2.5 or less. Metatibia inner spur length/metabasitarsus length: 0.4–0.5. Anteromesoscutum: mostly smooth or with shallow sparse punctures, except for anterior 0.3 where it has deeper and/or denser punctures. Mesoscutellar disc: mostly smooth. Number of pits in scutoscutellar sulcus: 11 or 12. Maximum height of mesoscutellum lunules/maximum height of lateral face of mesoscutellum: 0.2–0.3. Propodeum areola: partially defined by carinae on posterior 0.3–0.5 of its length, widely open anteriorly. Propodeum background sculpture: mostly sculptured. Mediotergite 1 length/width at posterior margin: 2.3–2.5. Mediotergite 1 shape: slightly widening from anterior margin to 0.7–0.8 mediotergite length (where maximum width is reached), then narrowing towards posterior margin. Mediotergite 1 sculpture: with some sculpture near lateral margins and/or posterior 0.2–0.4 of mediotergite. Mediotergite 2 width at posterior margin/length: 2.8–3.1. Mediotergite 2 sculpture: mostly smooth. Outer margin of hypopygium: inflexible (without a folded, transparent, semi–desclerotized area); with no pleats visible. Ovipositor thickness: anterior width at most 2.0 × posterior width (beyond ovipositor constriction) (?). Ovipositor sheaths length/metatibial length: 0.4–0.5. Length of fore wing veins r/2RS: 1.4–1.6. Length of fore wing veins 2RS/2M: 0.8 or less. Length of fore wing veins 2M/(RS+M)b: 1.1–1.3. Pterostigma length/width: 3.1–3.5. Point of insertion of vein r in pterostigma: about half way point length of pterostigma. Angle of vein r with fore wing anterior margin: more or less perpendicular to fore wing margin. Shape of junction of veins r and 2RS in fore wing: strongly angulated, sometimes with a knob.

Male. As in female.

Molecular data.

Sequences in BOLD: 6, barcode compliant sequences: 6.

Biology/ecology.

Gregarious (Fig. 257). Hosts: Tortricidae, Aesiocopa necrofolia.

Distribution.

Costa Rica, ACG.

Etymology.

We dedicate this species to Ana Piedra in recognition of her diligent efforts for the ACG Programa de Educacion Biológica.

Comments.

Apanteles anapiedrae shares with the diatraeae and guadaluperodriguezae groups a somewhat depressed body (dorso-ventrally), short antenna, and relatively small body size; however, it has an inflexible (unfolded) hypopygium without any pleats, a very small smooth area on lateral face of scutellum (0.2 × as high as maximum height of lateral face), and parasitizes a completely different group of Lepidoptera. The sculpture of propodeum and the areola shape are similar to species of the diatraeae group (but the latter group has a pleated hypopygium, a longer ovipositor, and the smooth area on lateral face of scutellum is at least 0.5 × as high as maximum height of lateral face). Apanteles anapiedrae does not resemble typical species of Apanteles because of its propodeal areola and unpleated hypopygium. It is likely to represent a derived species-group within Apanteles, or it might be placed in another genus. Pending further study of worldwide genera of Microgastrinae, we decided to describe the species under Apanteles because is the closest match at the moment.

Type locality.

COSTA RICA, Guanacaste, ACG, Sector Santa Rosa, Bosque San Emilio, 300m, 10.84389, -85.61384.

Holotype.

♀ in CNC. Specimen labels: 1. DHJPAR0013054. 2. 24 Apr. 2000, San Emilio Trap.

Description.

Female. Body color: body mostly dark except for some sternites which may be pale. Antenna color: scape, pedicel, and flagellum dark. Coxae color (pro-, meso-, metacoxa): dark, dark, dark. Femora color (pro-, meso-, metafemur): anteriorly dark/posteriorly pale, dark, dark. Tibiae color (pro-, meso-, metatibia): pale, pale, anteriorly pale/posteriorly dark. Tegula and humeral complex color: both dark. Pterostigma color: mostly pale and/or transparent, with thin dark borders. Fore wing veins color: partially pigmented (a few veins may be dark but most are pale). Antenna length/body length: antenna shorter than body (head to apex of metasoma), not extending beyond anterior 0.7 metasoma length. Body in lateral view: not distinctly flattened dorso–ventrally. Body length (head to apex of metasoma): 2.0 mm or less. Fore wing length: 2.0 mm or less. Ocular–ocellar line/posterior ocellus diameter: 2.0–2.2. Interocellar distance/posterior ocellus diameter: 1.7–1.9. Antennal flagellomerus 2 length/width: 2.6–2.8. Antennal flagellomerus 14 length/width: 1.1–1.3. Length of flagellomerus 2/length of flagellomerus 14: 2.0–2.2. Tarsal claws: simple. Metafemur length/width: 2.8–2.9. Metatibia inner spur length/metabasitarsus length: 0.4–0.5. Anteromesoscutum: mostly with deep, dense punctures (separated by less than 2.0 × its maximum diameter). Mesoscutellar disc: mostly smooth. Number of pits in scutoscutellar sulcus: 11 or 12. Maximum height of mesoscutellum lunules/maximum height of lateral face of mesoscutellum: 0.6–0.7. Propodeum areola: completely defined by carinae, including transverse carina extending to spiracle. Propodeum background sculpture: partly sculptured, especially on anterior 0.5. Mediotergite 1 length/width at posterior margin: 1.1–1.3. Mediotergite 1 shape: slightly widening from anterior margin to 0.7–0.8 mediotergite length (where maximum width is reached), then narrowing towards posterior margin. Mediotergite 1 sculpture: mostly sculptured, excavated area centrally with transverse striation inside and/or a polished knob centrally on posterior margin of mediotergite. Mediotergite 2 width at posterior margin/length: 4.4–4.7. Mediotergite 2 sculpture: with some sculpture, mostly near posterior margin. Outer margin of hypopygium: with a wide, medially folded, transparent, semi–desclerotized area; usually with 4 or more pleats. Ovipositor thickness: about same width throughout its length. Ovipositor sheaths length/metatibial length: 1.0–1.1. Length of fore wing veins r/2RS: 1.0 or less. Length of fore wing veins 2RS/2M: 1.4–1.6. Length of fore wing veins 2M/(RS+M)b: 0.9–1.0. Pterostigma length/width: 2.6–3.0. Point of insertion of vein r in pterostigma: about half way point length of pterostigma. Angle of vein r with fore wing anterior margin: more or less perpendicular to fore wing margin. Shape of junction of veins r and 2RS in fore wing: distinctly but not strongly angled.

Male. Unknown.

Molecular data.

Sequences in BOLD: 1, barcode compliant sequences: 1.

Biology/ecology.

Malaise trapped.

Distribution.

Costa Rica, ACG.

Etymology.

We dedicate this species to Ana Arias in recognition of her diligent efforts for the ACG Comedor in Santa Rosa.

Apanteles andreacalvoae Fernández-Triana, sp. n.

http://zoobank.org/176A4F60-5DB6-46F3-B83C-185F123842FB

http://species-id.net/wiki/Apanteles_andreacalvoae

Figs 68, 258
Apanteles Rodriguez10 (Smith et al. 2006). Interim name provided by the authors.
Type locality.

COSTA RICA, Guanacaste, ACG, Sector Cacao, Estación Gongora, 570m, 10.88700, -85.47443.

Holotype.

♀ in CNC. Specimen labels: 1. COSTA RICA, Guanacaste, ACG, Sector Cacao, Estación Gongora, 14.vi.2004, 570m, 10.88700, -85.47443, 04-SRNP-45868. 2. DHJPAR0001602.

Paratypes.

20 ♀, 6 ♂ (BMNH, CNC, INBIO, INHS, NMNH). COSTA RICA: ACG database codes: 04-SRNP-33636, 04-SRNP-45868, 09-SRNP-31397, DHJPAR0001605, DHJPAR0013013, DHJPAR0035406.

Description.

Female. Body color: body mostly dark except for some sternites which may be pale. Antenna color: scape, pedicel, and flagellum dark. Coxae color (pro-, meso-, metacoxa): dark, dark, dark. Femora color (pro-, meso-, metafemur): anteriorly dark/posteriorly pale, dark, dark. Tibiae color (pro-, meso-, metatibia): pale, pale, mostly pale but with posterior 0.2 or less dark. Tegula and humeral complex color: both dark. Pterostigma color: dark. Fore wing veins color: mostly dark (a few veins may be unpigmented). Antenna length/body length: antenna about as long as body (head to apex of metasoma); if slightly shorter, at least extending beyond anterior 0.7 metasoma length. Body in lateral view: not distinctly flattened dorso–ventrally. Body length (head to apex of metasoma): 2.0 mm or less, 2.5–2.6 mm, rarely 2.1–2.2 mm. Fore wing length: 2.1–2.2 mm or 2.7–2.8 mm. Ocular–ocellar line/posterior ocellus diameter: 2.0–2.2. Interocellar distance/posterior ocellus diameter: 1.7–1.9. Antennal flagellomerus 2 length/width: 3.2 or more. Antennal flagellomerus 14 length/width: 2.0–2.2. Length of flagellomerus 2/length of flagellomerus 14: 2.0–2.2. Tarsal claws: with single basal spine–like seta. Metafemur length/width: 3.2–3.3. Metatibia inner spur length/metabasitarsus length: 0.6–0.7. Anteromesoscutum: mostly with deep, dense punctures (separated by less than 2.0 × its maximum diameter). Mesoscutellar disc: mostly smooth. Number of pits in scutoscutellar sulcus: 7 or 8. Maximum height of mesoscutellum lunules/maximum height of lateral face of mesoscutellum: 0.6–0.7. Propodeum areola: completely defined by carinae, including transverse carina extending to spiracle. Propodeum background sculpture: partly sculptured, especially on anterior 0.5. Mediotergite 1 length/width at posterior margin: 2.9–3.1. Mediotergite 1 shape: more or less parallel–sided. Mediotergite 1 sculpture: with some sculpture near lateral margins and/or posterior 0.2–0.4 of mediotergite. Mediotergite 2 width at posterior margin/length: 2.8–3.1. Mediotergite 2 sculpture: mostly smooth. Outer margin of hypopygium: inflexible (without a folded, transparent, semi–desclerotized area); with no pleats visible. Ovipositor thickness: anterior width at most 2.0 × posterior width (beyond ovipositor constriction). Ovipositor sheaths length/metatibial length: 0.8–0.9. Length of fore wing veins r/2RS: 2.0–2.2. Length of fore wing veins 2RS/2M: 1.1–1.3. Length of fore wing veins 2M/(RS+M)b: 0.9–1.0. Pterostigma length/width: 3.6 or more. Point of insertion of vein r in pterostigma: about half way point length of pterostigma. Angle of vein r with fore wing anterior margin: more or less perpendicular to fore wing margin. Shape of junction of veins r and 2RS in fore wing: distinctly but not strongly angled.

Male. Similar to female.

Molecular data.

Sequences in BOLD: 7, barcode compliant sequences: 6.

Biology/ecology.

Gregarious (Fig. 258). Hosts: Hesperiidae