A revision of the parasitoid wasp genus Dolichogenidea Viereck (Hymenoptera, Braconidae) in the Neotropical region, with the description of 102 new species

Jose L. Fernandez-Triana; Caroline Boudreault; James B. Whitfield; Amelie Höcherl; M. Alex Smith; Winnifred Hallwachs; Daniel H. Janzen

doi:10.3897/zookeys.1237.141007

Monograph

A revision of the parasitoid wasp genus Dolichogenidea Viereck (Hymenoptera, Braconidae) in the Neotropical region, with the description of 102 new species

Jose L. Fernandez-Triana^‡, Caroline Boudreault^‡, James B. Whitfield^§, Amelie Höcherl^|, M. Alex Smith^¶, Winnifred Hallwachs^#, Daniel H. Janzen^#

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

§ University of Illinois, Urbana, United States of America

| Stuttgart State Museum of Natural History, Stuttgart, Germany

¶ University of Guelph, Guelph, Canada

# University of Pennsylvania, Philadelphia, United States of America

Corresponding author: Jose L. Fernandez-Triana ( cnc.braconidae@gmail.com )

Academic editor: Erinn Fagan-Jeffries

This is an open access article distributed under the terms of the CC0 Public Domain Dedication.

Citation: Fernandez-Triana JL, Boudreault C, Whitfield JB, Höcherl A, Smith MA, Hallwachs W, Janzen DH (2025) A revision of the parasitoid wasp genus Dolichogenidea Viereck (Hymenoptera, Braconidae) in the Neotropical region, with the description of 102 new species. ZooKeys 1237: 1-250. https://doi.org/10.3897/zookeys.1237.141007

ZooBank: urn:lsid:zoobank.org:pub:F77FA5A7-28CC-44B4-9428-D799119E4A18

Abstract

The parasitoid wasp genus Dolichogenidea is currently the second most speciose within the subfamily Microgastrinae (Hymenoptera: Braconidae), with 366 world species known so far, but with hundreds awaiting to be described. Here, the fauna of the Neotropical region is revised, with an emphasis in the Area de Conservación Guanacaste (ACG), Costa Rica. In addition to 23 species previously recorded from the Neotropics, 102 additional species are described as new, increasing the regional and world richness to 125 and 468 species, respectively. All species are diagnosed and described by using a combination of basic morphology (dichotomous key and brief diagnostic descriptions) and, when available DNA COI barcodes, biology (host data and wasp cocoon strategy), and distribution data. Neither morphology, biology, nor molecular data alone were sufficient to unambiguously separate all taxa, as all approaches were found to have limitations, but the combination of all three approaches provided stronger support to species delimitation. Morphology allowed the inclusion of all known species, therefore building a foundation upon which to improve as more molecular and biological data become available and new species are discovered; however, it was not sufficient (or it was very difficult to use) to separate at least 15% of all species keyed out in the dichotomous key. DNA barcoding was better able to separate species, and it is likely to become the most efficient way to identify species in the near future; however, DNA failed to identify 8.3% of the species with molecular data available, in addition to one third of the described species currently lacking molecular data. Biological data is currently the most incomplete, with only 42% of the species having associated host information, with a strong data availability bias towards ACG specimens. A total of 11 Lepidoptera families are here recorded to be parasitized by Neotropical Dolichogenidea, mainly Depressariidae (34% of all host data available), Gelechiidae (17%), Crambidae (14%), Tortricidae (10%), Thyrididae (8%) and Pyralidae (7%). Most of the wasps seem to be monophagous or at most oligophagous, as 56% are known to only parasitize a single host species, whereas 23% parasitize two host species and 10% parasitize three hosts; in almost all cases, the hosts species belong to one genus (or related genera) in the same Lepidoptera family. Most species of Dolichogenidea are found between 400–1,500 m, but a few have been found at higher elevations, including a few examples higher than 3,000 m (Costa Rica) and 4,000–4,100 m in the Andes (South America). The following nomenclatural acts are proposed: 1) the genus Exoryza is synonymized under Dolichogenidea, syn. nov.; 2) a total of 16 species are transferred to Dolichogenidea as comb. nov., one species formerly in the genus Apanteles: Dolichogenidea croceicornis (Muesebeck, 1958) and all 15 species formerly placed within Exoryza (six of them from the Neotropics): Dolichogenidea asotae (Watanabe, 1932), Dolichogenidea belippicola (Liu & You, 1988), Dolichogenidea hylas (Wilkinson, 1932), Dolichogenidea mariabustosae (Fernandez-Triana, 2016), Dolichogenidea megagaster (de Saeger, 1944), Dolichogenidea minnesota (Mason, 1981), Dolichogenidea monocavus (Valerio & Whitfield, 2004), Dolichogenidea oryzae Walker, 1994, Dolichogenidea reticarina (Song & Chen, 2003), Dolichogenidea richardashleyi (Fernandez-Triana, 2016), Dolichogenidea ritaashleyae (Fernandez-Triana, 2016), Dolichogenidea rosamatarritae (Fernandez-Triana, 2016), Dolichogenidea safranum (Rousse & Gupta, 2013), Dolichogenidea schoenobii (Wilkinson, 1932) and Dolichogenidea yeimycedenoae (Fernandez-Triana, 2016); 3) Dolichogenidea yeimycedenoae (Fernandez-Triana, 2016) becomes a senior secondary homonym of Dolichogenidea yeimycedenoae Fernandez-Triana & Boudreault, 2019; therefore, Dolichogenidea cedenoae Fernandez-Triana & Boudreault, 2025 is a replacement name for Dolichogenidea yeimycedenoae Fernandez-Triana & Boudreault, 2019; 4) the following 102 species, all authored by Fernandez-Triana & Boudreault, are described as sp. nov.: D. aceituno, D. alanflemingi, D. alejandromarini, D. alerce, D. alexamasisae, D. alexandrei, D. alixhamiltonae, D. amazonas, D. anacamposae, D. andreamezae, D. angelsolisi, D. anikenpalolae, D. anniapicadoae, D. annlisterudae, D. annychaverae, D. antioquia, D. antjevirkusae, D. arenal, D. bernardoespinozai, D. beryllacosteae, D. bradzlotnicki, D. caldas, D. carlosalvaradoi, D. carlosviquezi, D. chichicastenango, D. christinaagapakisae, D. claudiadoblesae, D. dole, D. encruzilhada, D. ericpalolai, D. ericsimoni, D. escobarae, D. felipechavarriai, D. frankjoycei, D. fredhicksi, D. helenedumasae, D. heredia, D. ingredolsonae, D. isabelleae, D. isidrochaconi, D. jaimelewisi, D. jasonkelleyi, D. jennyphillipsae, D. jessiehillae, D. johnrobinsoni, D. jorgecarvajali, D. jorgecortesi, D. josephfridmani, D. joshdarfleri, D. juanmatai, D. junhyongkimi, D. kasiiya, D. katiemccluskeyae, D. kenzabaddouae, D. lacochaparamo, D. leahdennisae, D. limoncocha, D. luishamiltoni, D. luzmariaromeroae, D. machupichu, D. mehdirheljari, D. moniqueae, D. moniquegilbertae, D. ninamasisae, D. nothofagus, D. oiketicus, D. palenque, D. papallacta, D. paulfryi, D. pedroleoni, D. puschendorfi, D. putumayo, D. puyo, D. rexhamiltoni, D. robertofernandezi, D. robinsherwoodae, D. robmacewani, D. robpringlei, D. rociocordobae, D. rodrigogamezi, D. ronaldzunigai, D. rubymacpearsae, D. rudyamadori, D. sallydaleyae, D. sarahoconnorae, D. scottmilleri, D. shelleymcsweeneyae, D. sigifredomarini, D. stephmae, D. stevestroudi, D. susanabramsae, D. teremariae, D. tiboshartae, D. timrichi, D. tomdaleyi, D. tristanpalolai, D. tucuman, D. verobrondexae, D. virgendelparamo, D. weaversway, D. yungas, D. yvesbraeti.

Key words:

DNA barcoding, host data, Microgastrinae, morphology, Neotropics, turbo taxonomy

Introduction

With 366 described species, the genus Dolichogenidea Viereck, 1911 is the second most speciose taxon within Microgastrinae parasitoid wasps (Hymenoptera: Braconidae), only surpassed by Apanteles (Fernandez-Triana et al. 2020). Dolichogenidea was originally described as a subgenus of Apanteles, but it was later elevated to generic rank by Mason (1981). Since then, its recognition has been at times controversial (e.g., Mason 1981; van Achterberg 2003; Fernandez-Triana et al. 2014a, 2020), but in the past decade it has been widely accepted as a valid genus by most of the taxonomists working with the group worldwide.

This paper aims to describe a significant number of new species from the Neotropical region. We provide a key, illustrations for all known species, and comments on natural history and DNA barcoding data. Special emphasis is given to the Area de Conservación de Guanacaste, northwestern Costa Rica (ACG), as the present study contributes taxonomically to its ongoing insect biodiversity inventory based on rearing and Malaise trapping (Janzen and Hallwachs 2016). Other areas beyond ACG are also dealt with, although they have not been as comprehensively studied.

Materials and methods

In this paper we diagnose and describe all the species of Dolichogenidea known from the Neotropical region by using a combination of basic morphology (dichotomous key and brief diagnostic descriptions), DNA barcodes (COI gene, when available), biology (host data and wasp cocoon strategy, when available), and distribution data, following suggestions outlined in Fernandez-Triana (2022).

Morphological terms and measurements mostly follow Huber and Sharkey (1993), Whitfield (1997), Karlsson and Ronquist (2012) and Fernandez-Triana et al. (2014a). The abbreviations T1, T2, and T3 are used for metasomal mediotergites 1, 2, and 3 respectively, and F1–16 refer to flagellomeres 1 to 16.

Species are diagnosed to species groups following some simple characters and then within each group a few additional characters are provided to separate species in that specific group. The resulting diagnostic descriptions are thus based on a relatively small set of morphological characters which vary depending on the species group.

Complete and verbatim label details are provided only for holotypes. Paratypes and other specimens examined are listed only with basic information (country, repository, sex, and voucher codes). All information associated with those specimens can be accessed in the publicly available CNC database (https://www.cnc.agr.gc.ca/taxonomy/TaxonMain.php). A dataset with all details for specimens with available DNA COI sequences in BOLD is available at https://dx.doi.org/10.5883/DS-NEODOLIC.

DNA barcoding was also used to characterize and recognize species. DNA extracts were obtained from single legs using a glass fiber protocol (Ivanova et al. 2006), and total genomic DNA was re-suspended in 30 μl of distilled water. The barcode region, a 658 base pairs (bp) region near the 5’ terminus of the COI gene, was amplified using standard primers following established protocols (e.g., see references in Fernandez-Triana et al. 2014a).

The Barcode Index Number (BIN) was considered to approximately characterize species limits, following the BIN concept detailed in Ratnasingham and Hebert (2013). A Neighbor Joining (NJ) tree with all available sequences of Neotropical Dolichogenidea (Suppl. material 1) and with single sequences per BIN were generated using BOLD capabilities.

For additional analyses, best-fit models were identified using ModelFinder (Kalyaanamoorthy et al. 2017), Maximum Likelihood analysis was performed using IQ-TREE 2 (Minh et al. 2020) including ultrafast Bootstrap (Hoang et al. 2018). Bayesian analysis was performed using BEAST X v. 1.10.4 (Suchard et al. 2018) with BEAGLE (Ayres et al. 2012). For those BINs that showed potential BIN-discrepancies, TCS haplotype network analysis was performed using PopART (Snell et al. 2002; Leigh and Bryant 2015). The sequences used were selected based on sequence length which is indicated in the description of the figures depicting the haplotype networks. In the figures, each hatch mark in the network represents a single mutational change; small black dots at nodes indicate missing haplotypes. The diameter of the circles is proportional to the number of specimens sampled per haplotype. We also performed clustering using ASAP (Puillandre et al. 2021) for some selected species

All information for the sequences associated with each specimen barcoded (including primers and trace files) is available on the Barcode of Life Data System (BOLD) (Ratnasingham and Hebert 2007) under the dataset “DS-NEODOLI Neotropical Dolichogenidea species” (https://www.boldsystems.org/index.php/MAS_Management_DataConsole?codes=DS-NEODOLI).

Host data (Lepidoptera species) as well as wasp cocoon strategy (solitary/gregarious) were mostly taken from the website “Dynamic database for an inventory of the macrocaterpillar fauna, and its food plants and parasitoids in ACG databases” (http://janzen.sas.upenn.edu/caterpillars/database.lasso). We caution that considering a wasp species “solitary” when there are only one or two rearing records may be an artifact of their having been just one surviving wasp larva of a gregarious species; e.g., see record DHJPAR0042935, for wasp species Dolichogenidea alejandromasisi, where there is just a single cocoon of a species that is notoriously gregarious (https://bench.boldsystems.org/index.php/MAS_DataRetrieval_OpenSpecimen?selectedrecordid=ASHYH693-11).

Although many hosts are identified to species level in that database, for many others the available information only included the genus of Lepidoptera with an interim or provisional species name code (supported by biology or DNA barcoding to be one biological unit); these conventions to record Lepidoptera hosts from ACG have been used in many scientific papers published during the past 10+ years (see Fernandez-Triana et al. 2023 for an explanation on how to interpret those codes).

We also note that, in contrast to parasitoids of macrolepidopterans (e.g., Nymphalidae, Saturniidae, Sphingidae, Geometridae) whose larvae are distinctive and easy to identify in the field, many hosts of Dolichogenidea have small and poorly identified larvae often hidden in leaf rolls or nests and therefore subject to misidentification other than when there are so many that some survive to adult status, thereby ensuring the host identification (e.g., 93 rearings of Dolichochenidea alejandromasisi from the single species Antaeotricha renselariana (Depressariidae) eating 32 species of Fabaceae in ten genera in all ACG habitats).

The majority of the photos were taken with a Keyence VHX-1000 Digital Microscope (Keyence Corporation, Japan), using a lens with a range of 10–130×; multiple images were taken of the structures through the focal plane and then combined to produce a single in-focus image using the software associated with the Keyence System. Some photographs were taken with a Canon EOS-7D Mark 2 (G) (Canon Inc., Japan) using a super-macro lens Canon MP-65 with a Yongnuo professional flash speedlight flashlight installed on a modified microscope stand; multiple images (in raw format .CR2) were taken of a structure through the focal plane, converted to .dng with Adobe DNG converter, then corrected (brightness and contrast) in Adobe Bridge CS4, converted to .tiff images with Adobe Photoshop CS4 and finally combined to produce a single in-focus image using Zerene Stacker (http://zerenesystems.com/cms/stacker). Final images were corrected using GIMP 2.10.12. Images of wasp cocoons and larvae were taken by parataxonomists at ACG with a Canon camera. All plates were prepared using Microsoft PowerPoint 2010 and saved as .TIF files.

Images of one holotype (of a previously described species), deposited in the Natural History Museum (Smithsonian Institution, Washington DC, USA) were accessed through the Primary Type Specimens Catalog of the Department of Entomology Collections (https://collections.nmnh.si.edu/search/ento/). The downloaded images were later combined into plates. Those images were classified in that website as CC0, therefore making them available under the Creative Commons license CC0 1.0 license granting the right to share for personal and educational purposes under the fair use doctrine; in any case, we acknowledge the source of those images here.

All ACG specimens were collected, exported, and DNA barcoded under Costa Rican government permits issued to BioAlfa (R-054-2022-OT-CONAGEBIO; R-019-2019-CONAGEBIO; National Published Decree #41767), JICA-SAPI #0328497 (2014) and DHJ and WH (ACG-PI-036-2013; R-SINAC-ACG-PI-061-2021; Resolución N°001-2004 SINAC; PI-028-2021).

Results

This paper deals with 127 species (Table 1). This total includes 125 Neotropical species (102 of them described as new) as well as the only two Nearctic species of Dolichogenidea recorded from southern areas in the United States around the Gulf of Mexico (Alabama, Florida, Louisiana, Mississippi, and Texas): D. acrobasidis (Muesebeck, 1921) and D. bushnelli (Muesebeck, 1933). The Nearctic species were included in the key below in case they are later found to also occur in the Neotropics, especially Mexico or on some Caribbean islands, although at present there is no evidence they occur there.

Table 1.

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List of all known Neotropical Dolichogenidea species with associated molecular data (BIN code, when available, or details of partial barcodes), host families and host species, wasp cocoon strategy (S- solitary; G- gregarious), and known distribution by country. [See explanation in Materials and methods about few cases where considering a wasp cocoon as “solitary” may not be accurate].

Species name	Molecular data	Host Family	Host species	S/G	Distribution
Dolichogenidea aceituno					Chile
D. acrobasidis		Pyralidae, Tortricidae	Acrobasis caryae, Gretchena bolliana	S	United States
D. alanflemingi	BOLD:AAN2497	Depressariidae	Antaeotricha Janzen49, Antaeotricha Janzen146	S	Costa Rica, Saint Vincent, Trinidad & Tobago
D. alejandromarini	BOLD:AAI9746				Costa Rica
D. alejandromasisi	BOLD:ABX6174	Depressariidae	Antaeotricha renselariana	G	Costa Rica
D. alerce	BOLD:AAH1316				Chile
D. alexamasisae	BOLD:AAF5364	Crambidae	Herpetogramma Janzen04, Rhectocraspeda Solis05	S	Costa Rica, Ecuador, Venezuela
D. alexandrei					Guatemala
D. alixhamiltonae	BOLD:AAL2285	Thyrididae	Banisia myrsusalisDHJ01	S	Costa Rica
D. amazonas					Peru
D. anacamposae	BOLD:AAE8612	Tortricidae	Olethreutes Brown22, Olethreutes Janzen323	S	Costa Rica
D. andreamezae	BOLD:ABA7252	Erebidae	Rivula Poole03	S	Costa Rica
D. angelagonzalezae	BOLD:AAL2298	Choreutidae	Brenthia Janzen12	S	Costa Rica
D. angelsolisi	BOLD:ACI3413	Immidae	immidJanzen01 Janzen26	S	Costa Rica
D. anikenpalolae	BOLD:ABY1812	Crambidae	spiloBioLep01 BioLep379	S	Costa Rica
D. anniapicadoae	BOLD:ABY7999	Crambidae	Ategumia lotanalis	S	Costa Rica
D. annlisterudae	BOLD:AAB5549, BOLD:ACF0272				Costa Rica
D. annychaverae	BOLD:AAD5258				Costa Rica
D. antioquia					Colombia
D. antjevirkusae	BOLD:AAM5849				Costa Rica
D. arenal					Costa Rica
D. bernardoespinozai	BOLD:AAE8596				Costa Rica
D. beryllacosteae	BOLD:AAM1098	Thyrididae	Microsca hedialis	G	Costa Rica
D. bradzlotnicki	BOLD:ACC1295	Depressariidae	Chlamydastis vividella, Stenoma Janzen199	S	Costa Rica
D. bushnelli		Pyralidae, Tortricidae	Dioryctria abietella, Dioryctria clarioralis, Dioryctria disclusa, Rhyacionia bushnelli, Rhyacionia pasadenana	S	United States
D. caldas					Colombia
D. carlosalvaradoi	BOLD:AAM5848				Costa Rica
D. carlosmanuelrodriguezi	BOLD:ABZ4155	Depressariidae	Antaeotricha spurca	S	Costa Rica
D. carlosviquezi					Costa Rica
D. cedenoae	BOLD:ABY3724	Depressariidae	elachJanzen01 Janzen196, elachJanzen01 Janzen397, Antaeotricha Janzen126	G	Costa Rica
D. chichicastenango					Guatemala
D. christinaagapakisae	BOLD:AAI9755	Depressariidae, Gelechiidae	Gonionota Janzen22, gelJanzen01 Janzen23	S	Costa Rica
D. claudiadoblesae	BOLD:AAD2236				Costa Rica
D. croceicornis		Crambidae	Microthyris anormalis	S	Peru
D. dole	BOLD:AAM5739				Costa Rica
D. encruzilhada					Brazil
D. ensiger	BOLD:AAA3764	Crambidae, Tortricidae	Fissicrambus mutabilis, Neodactria zeellus, Choristoneura freemani, Epiblema strenuana	?S	Canada, Costa Rica, United States
D. ericpalolai	BOLD:AAF7717				Costa Rica
D. ericsimoni					Chile
D. escobarae					Brazil
D. evadne					Juan Fernández Islands
D. felipechavarriai	BOLD:ACC4119	Depressariidae	Gonionota Janzen116	S	Costa Rica
D. frankjoycei	BOLD:ABA3469	Tortricidae	Platynota rostrana DHJ01, Platynota rostrana DHJ02	S	Costa Rica
D. fredhicksi	BOLD:AAK2061	Depressariidae	Anadasmus Janzen25, Stenoma Janzen27	G	Costa Rica
D. gelechiidivoris	BOLD:AAM4042	Gelechiidae	Phthorimaea operculella, Keiferia lycopersicella, Tuta absoluta	S	Algeria, Chile, Colombia, Peru, Spain, Venezuela
D. genuarnunezi	BOLD:ACC1300	Depressariidae	Antaeotricha phaeoneura	G	Costa Rica
D. hedyleptae		Crambidae	Maruca vitrata, Omiodes indicata	?G	Puerto Rico
D. helenedumasae	BOLD:ABZ4155				Brazil, French Guiana
D. heredia					Costa Rica
D. homoeosomae	Partial sequences (425 bp)	Pyralidae	Homoeosoma electellum	S	Cuba, Canada, United States
D. ingredolsonae	BOLD:AAM5850				Costa Rica
D. isabelleae					Ecuador
D. isidrochaconi	BOLD:AAB9372				Costa Rica
D. jaimelewisi	BOLD:AAM5738	Crambidae	Herpetogramma salbialis, Herpetogramma Dapkey27, spiloBioLep01 BioLep617	G	Costa Rica
D. jasonkelleyi	BOLD:AAI9747				Costa Rica
D. jennyphillipsae	BOLD:AAM5088				Costa Rica
D. jessiehillae	BOLD:AAM5851				Costa Rica
D. johnrobinsoni	BOLD:ACF0267				Costa Rica
D. jorgecarvajali	BOLD:AAM5847				Costa Rica
D. jorgecortesi	BOLD:AAM5846				Costa Rica
D. josealfredohernandezi	BOLD:ABA9255	Depressariidae	Stenoma Janzen99	S	Costa Rica
D. josephfridmani	BOLD:AAE8602				Costa Rica
D. joshdarfleri	BOLD:AAC7481				Costa Rica
D. juanmatai	BOLD:AAM5740				Costa Rica
D. junhyongkimi	BOLD:AAD6850				Costa Rica
D. kasiiya	BOLD:AAM5750				Costa Rica
D. katiemccluskeyae	BOLD:ACE8228				Costa Rica, Honduras
D. kenzabaddouae	BOLD:AAY4695	Depressariidae	Antaeotricha Janzen221	S	Costa Rica
D. lacochaparamo					Colombia
D. leahdennisae	BOLD:AAJ1396				Costa Rica
D. limoncocha					Ecuador
D. luishamiltoni	BOLD:AAT8840	Gelechiidae	gelJanzen01 Janzen394	S	Costa Rica
D. luzmariaromeroae	BOLD:ABX5620	Pyralidae	phyBioLep01 BioLep758	S	Costa Rica
D. machupichu					Peru
D. mariabustosae	BOLD:AAN2496	Gelechiidae	gelJanzen01 Janzen319	S	Costa Rica
D. mehdirheljari	BOLD:AAM5852				Costa Rica
D. melaniamunozae	BOLD:AAB5701	Depressariidae	Cerconota recurvella	S	Costa Rica
D. moniqueae					Chile
D. moniquegilbertae	BOLD:AAX8653				Costa Rica, Mexico
D. monocavus					Costa Rica
D. ninamasisae	BOLD:AAY4690	Tortricidae	Megalota crassana, Megalota spinulosa	S	Costa Rica
D. nothofagus					Chile
D. oiketicus		Psychidae	Oiketicus kirbyi	S	Trinidad & Tobago
D. palenque					Ecuador
D. papallacta					Ecuador
D. parallelis					Saint Vincent
D. paulfryi	BOLD:ACF2929				Costa Rica
D. pedroleoni	BOLD:AAB4946	Mimallonidae	Eadmuna Janzen01	G	Costa Rica
D. phthorimaeae		Gelechiidae	Keiferia glochinella, Keiferia inconspicuella, Keiferia lycopersicella	S	Canada, Honduras, United States
D. politiventris					Colombia, Dominican Republic, Puerto Rico, Saint Vincent, Trinidad & Tobago
D. puschendorfi	BOLD:AAM5853				Costa Rica
D. putumayo					Colombia
D. puyo					Ecuador
D. rexhamiltoni	BOLD:AAL2287				Costa Rica
D. richardashleyi	BOLD:ABX6267	Gelechiidae	gelJanzen01 Janzen349	S	Costa Rica
D. ritaashleyae	BOLD:ABX6267				Costa Rica
D. robertofernandezi	BOLD:AAC8392				Costa Rica
D. robinsherwoodae	BOLD:ABX5195	Depressariidae	Antaeotricha Janzen221	S	Costa Rica
D. robmacewani					Brazil
D. robpringlei		Thyrididae	Collinsa ferreiceps	G	Costa Rica
D. rociocordobae	BOLD:ACJ2777	Gelechiidae	Dichomeris Janzen273, Dichomeris Janzen703	S	Costa Rica
D. rodrigogamezi	BOLD:AAM5843				Costa Rica
D. rogerblancoi	BOLD:AAL2325	Depressariidae	Antaeotricha radicalisDHJ01	G	Costa Rica
D. ronaldzunigai	BOLD:AAT8860	Depressariidae	Chlamydastis montywoodi, Chlamydastis tryphon, Chlamydastis vividella, Stenoma Janzen199, elachJanzen01 Janzen693	S	Costa Rica
D. rosamatarritae	BOLD:ABY5258	Choreutidae, Gelechiidae, Depressariidae	Brenthia Janzen05, Stenoma Phillips543, gelJanzen01 Janzen16	S	Costa Rica
D. rubymacpearsae					Ecuador, Peru
D. rudyamadori	BOLD:AAX8664				Costa Rica
D. sallydaleyae	BOLD:ACM2280	Depressariidae	elachBioLep01 BioLep286	S	Costa Rica
D. sarahoconnorae	BOLD:ABX6008	Thyrididae	Microsca hedialis, Microsca polychloralis, siculoJanzen01 biolep03, siculoJanzen01 Janzen05	G	Costa Rica
D. scottmilleri	BOLD:AAC2174, BOLD:ACE8823	Thyrididae	Microsca paullula	G	Costa Rica
D. shelleymcsweeneyae	BOLD:AAM5736				Costa Rica
D. sigifredomarini	BOLD:ACI3397	Depressariidae		S	Costa Rica, French Guiana
D. stephmae					Brazil
D. stevestroudi	BOLD:ACB1629	Gelechiidae	gelJanzen01 Janzen22	S	Costa Rica
D. susanabramsae	BOLD:AAI6323	Gelechiidae	Dichomeris designatellaDHJ04	S	Costa Rica
D. teremariae	BOLD:AAM5842				Costa Rica, Honduras
D. tiboshartae	BOLD:AAC5949	Depressariidae, Gelechiidae	Dichomeris Janzen76, elachJanzen01 Janzen409, gelJanzen01 Janzen116	S	Costa Rica
D. timrichi	BOLD:AAJ1390				Costa Rica
D. tomdaleyi	BOLD:AAD8952	Depressariidae, Crambidae	Antaeotricha incrassata, Antaeotricha cirrhoxanthaDHJ02, Antaeotricha similisEPR01, Antaeotricha similisEPR02, Antaeotricha BioLep46, Antaeotricha Janzen23, Antaeotricha Janzen31, Antaeotricha Janzen77, Antaeotricha Janzen106, Antaeotricha Janzen146, Antaeotricha Janzen290, Antaeotricha Janzen292DHJ0, Antaeotricha Janzen364, Antaeotricha Philips01, Chlamydastis Janzen04, Stenoma Janzen18, Stenoma Janzen58, Stenoma Janzen199, Stenoma Janzen699	S	Costa Rica
D. tristanpalolai	BOLD:AAI9740				Costa Rica
D. tucuman					Argentina
D. verobrondexae					Venezuela
D. virgendelparamo					Ecuador
D. weaversway		Gelechiidae	Telphusa BioLep476		Costa Rica
D. yeimycedenoae	BOLD:ABX6267				Costa Rica
D. yungas					Bolivia
D. yvesbraeti					French Guiana

Of the 25 species previously described, seven had been previously placed in different genera but are here transferred to Dolichogenidea as new combinations: D. croceicornis (Muesebeck, 1958) which was described within Apanteles (based on holotype having vannal lobe entirely setose); and species formerly placed within Exoryza (see below for rationale to synonymize this genus within Dolichogenidea): D. mariabustosae (Fernandez-Triana, 2016), D. monocavus (Valerio & Whitfield, 2004), D. richardashleyi (Fernandez-Triana, 2016), D. ritaashleyae (Fernandez-Triana, 2016), D. rosamatarritae (Fernandez-Triana, 2016) and D. yeimycedenoae (Fernandez-Triana, 2016). A special situation arises with the species Exoryza yeimycedenoae Fernandez-Triana, 2016 which, when here transferred to Dolichogenidea yeimycedenoae (Fernandez-Triana, 2016), becomes a senior secondary homonym of Dolichogenidea yeimycedenoae Fernandez-Triana & Boudreault, 2019; because the latter name now becomes a junior secondary homonym, that species would need a replacement name. Therefore, here we propose Dolichogenidea cedenoae Fernandez-Triana & Boudreault, nom. nov. as a replacement name for Dolichogenidea yeimycedenoae Fernandez-Triana & Boudreault, 2019.

The status of Exoryza as a valid genus, separate from Dolichogenidea, has been questioned by many authors (Valerio et al. 2004; Rousse and Gupta 2013; Fernandez-Triana et al. 2014a, 2016, 2020). The distinction may be particularly difficult because many species of Dolichogenidea have T1 and T2 strongly sculptured and T2 with a similar shape as that described for Exoryza (Mason 1981). Molecular data (DNA barcodes) do not support the separation of these two genera, but retrieve Exoryza as one group within Dolichogenidea (e.g., Fig. 2; see also Suppl. material 1). Here we formally synonymize Exoryza under Dolichogenidea. This new synonymy affects the six Neotropical species mentioned above as well as nine other species from other biogeographical regions formerly placed within Exoryza which are here also transferred as new combinations: Dolichogenidea asotae (Watanabe, 1932), D. belippicola (Liu & You, 1988), D. hylas (Wilkinson, 1932), D. megagaster (de Saeger, 1944), D. minnesota (Mason, 1981), D. oryzae Walker, 1994, D. reticarina (Song & Chen, 2003), D. safranum (Rousse & Gupta, 2013), and D. schoenobii (Wilkinson, 1932).

A total of 73 species are here described as new from Costa Rica, 71 of them from ACG. When accounting for additional species previously described (Valerio et al. 2004; Fernandez-Triana et al. 2016, 2019), there are now 87 Dolichogenidea species recorded from Costa Rica, 84 of them from ACG alone.

The rest of the Neotropical region has barely been studied, as we record here only a few species from other countries, Ecuador (8 species), Chile (7, including one species from Juan Fernandez Islands), Brazil and Peru (5), French Guiana, Honduras, Saint Vincent, Trinidad & Tobago, and Venezuela (3), Guatemala and Puerto Rico (2), Argentina, Bolivia, Cuba, Dominican Republic, and Mexico (1). Therefore, the 124 Neotropical species of Dolichogenidea accounted for within this paper represent only a small fraction of the actual species richness in the region. We assume that there are hundreds of species remaining undescribed in collections and in nature, especially from South America.

Including the 102 new taxa described here, Dolichogenidea now comprises 468 species. The actual diversity of the genus was recently estimated to be at least 700 species (Fernandez-Triana et al. 2020) but this now seems like a rather severe underestimate.

Morphology and species concepts used in this paper

Rather few taxonomic publications specify the species concept used (Williams and Ebach 2020), therefore we would like to spend a bit of time explaining how the different data sources were used to generate species hypotheses for Neotropical Dolichogenidea and why it is important to incorporate as many sources of information as possible. We used an integrative taxonomy approach where species were recognized, delimited and diagnosed using a combination of morphological, molecular and biological traits – insofar as these data were available. Morphological data are included in a dichotomous key, species diagnostic descriptions, and comprehensive illustrations for all 127 species dealt with in this review. Molecular data included DNA barcodes (complete or partial sequences), which were available for 67.7% of all Neotropical species. Biological data included parasitoid strategy (solitary or gregarious) and host-parasitoid associations, which were available for 41.9% of all Neotropical species. Geographical distribution was not used as a criterion to separate species but it was informative, particularly in deciding which described species should be considered and whether species were conspecific with existing types (it may also be that two or more “conspecifics” are found later to be reliably distinguished by geography, e.g., upper elevations versus adjacent lowlands, as it has been the case with other ACG groups, e.g., Smith et al. 2023).

Morphological data, especially in the dichotomous key and species diagnoses, were intended to provide a foundation for dealing with and incorporating species described in the future. As more species are discovered and described, they can be added to the key, and some of the species diagnoses may need to be modified to accommodate additional species. Morphology also allows us to include all known species in our taxonomic framework, including those lacking molecular and biological data at present.

At the beginning of the work for this revision, a number of morphological characters were selected, and we started scoring them for several species. Some species were very similar morphologically, suggesting potential complexes of cryptic species, as has been the case with many other ACG sympatric species. Evaluating those with more detailed analyses (e.g., morphometrics) was not possible because most of the species were not represented by enough specimens to properly assess morphological variability. Additionally, some characters were difficult to measure properly (e.g., the length of T1, the width of the posterior margin of T1, the length of the setose part of the ovipositor sheath), and thus are liable to different interpretations, with resulting measurements not being accurate enough to be useful. Therefore, we decided to use a simplified approach, where species were first grouped based on characters which were simple to assess and score and then, within each of those groups, additional characters were used to recognize individual species. It must be stressed that many of the morphological groupings are not thought to be monophyletic but are only used as a way to organize the work of recognizing species. For example, just by assessing the shape and sculpture of T1 and T2 it is possible to separate all Neotropical species of Dolichogenidea in to three groups (Fig. 1A). The first group is characterized by having T1 mostly smooth and rectangular, and T2 smooth and quadrate to broadly rectangular; it includes nine species so far recorded from the Neotropics and it certainly appears to represent a monophyletic group, supported by morphology, DNA and biology (Fernandez-Triana et al. 2019). The second group includes 29 species characterized by having T1 and T2 strongly sculptured with longitudinal striae and with T2 subquadrate to rectangular (or, in a few cases, transverse); it can be divided further into two distinctive subgroups, one of them certainly monophyletic and comprising what until now was considered to be the genus Exoryza (Fernandez-Triana et al. 2016) while the other subgroup is probably not monophyletic but still recognizable because of the distinctive shape and relatively coarse sculpture of T1 and T2 (see details in the dichotomous key below). The third group is the largest with 87 species and it is almost certainly not monophyletic. The third group can be separate into two subgroups based on the color of the coxae (Fig. 1B). And these two subgroups can be separated further based on body color, leg color, T1 shape, ovipositor sheath length, or other unique characters (e.g., Fig. 1C, D). The resulting key uses a relatively small set of characters which separate several groups, and then all species within each group are further diagnosed by adding additional characters (which work at that level but may not work for another group).

Figure 1.

Simplified depiction of characters used to separate all Neotropical species of Dolichogenidea A separation of species in three main groups B further separation of one of the main groups in two subgroups C, D additional separation of some subgroups into smaller units. The characters shown are simplified and abbreviated, for complete details the dichotomous key must be consulted.

This “modular approach” considerably sped up the work of diagnosing and describing the taxa, but it also meant that the diagnostic descriptions provided do not include the same set of characters assessed for every species: they only include characters needed for a specific group. It can be argued that, when future studies find additional new species of Neotropical Dolichogenidea, it is likely that both the dichotomous key and the diagnostic descriptions will need to be revised, modified or expanded accordingly, perhaps in some cases significantly. But the same will happen when more DNA barcodes are obtained, or new host-parasitoid associations are discovered. Therefore, the present work must be taken as a first approximation, which can/should be improved as knowledge advances.

For 11 species (8.7%) morphology alone was insufficient or very subjective, and DNA barcodes were incorporated into the key as the only available or the most reliable way to separate two species (e.g., see couplets 5, 20, 21, 38, 64, and 68 in the key below). Additionally, for at least another eight species (6.3%) the morphological characters provided in the key are either difficult to assess or interpret and may not be distinctive enough to clearly and easily differentiate species (e.g., see couplets 9, 60, 77 and 86 in the key below). Therefore, morphology alone is not sufficient (or it is very difficult to use) for at least 15% of all species keyed out in the dichotomous key we provide.

Despite its shortcomings for species diagnosis, morphology allows us to include all Neotropical species of Dolichogenidea known at present into a common framework, building a foundation upon which to improve as more molecular and biological data become available and new species are discovered.

DNA barcoding and the molecular identification of Neotropical Dolichogenidea

Bayesian (Fig. 2; see also Suppl. material 1) and Maximum Likelihood (Suppl. material 2) analyses did not recover Dolichogenidea as monophyletic. While our analyses have the limitation of including only a single gene, more comprehensive molecular analyses (e.g., Parks 2018’s five-gene molecular phylogeny; ongoing research based on data from Jasso-Martinez et al. 2022’s ultra-conserved elements) have also found the genus to be polyphyletic. The difficulties to characterize Dolichogenidea, both morphologically and molecularly, and to separate it from putatively related genera (such as some Pholetesor, Apanteles, and Parapanteles) have long been recognized and extensively discussed (e.g., Fernandez-Triana et al. 2014a, 2016, 2020; Parks 2018). Regardless of the current challenges to unambiguously recognize the genus, most species with molecular data available can be reliably identified using DNA barcodes.

Figure 2.

Bayesian inference consensus topology of the COI barcoding region of the Dolichogenidea species dataset, including additional outgroups. Triangles indicate collapsed branches. Branch labels represent p-values. Pale blue coloration indicates species from other genera and outgroups, orange coloration indicates the cluster with species that were formerly part of Exoryza. The sequences and BEAST settings used can be retrieved from Suppl. material 1.

The current coverage of DNA barcodes for Neotropical Dolichogenidea is fairly comprehensive, with 68.5% of the species being represented in BOLD as of April 2024 by at least 83 BINs. A total of 84 species (67.7%) have barcode compliant sequences while another one, D. homoeosomae, has two almost complete sequences of 425 bp and it is also included in the discussion below.

While most species had a unique correspondence with a single BIN, seven cases included either multiple BINs per single species or more than one species within a BIN, they are briefly discussed below.

Dolichogenidea scottmilleri sequences are included in BINs BOLD:AAC2174 and BOLD:ACE8823 (Fig. 3A), with a minimum p-distance between them of 1.05% (5 bp). Specimens from both BINs were reared from what appears to be the same Lepidoptera hosts, and we could not find any morphological difference to separate them, therefore we consider these two BINs to represent the same species.

Figure 3.

A TCS haplotype network of BINs BOLD:AAC2174 and BOLD:ACE8823, sequence length for analysis: 561 bp. Host species are indicated via coloration B TCS haplotype network of BINs BOLD:AAB5549 and BOLD:ACF2930, sequence length for analysis: 626 bp. The different molecular clusters resulting from ASAP analysis are indicated via coloration.

Sequences of D. annlisterudae are included in BINs BOLD:AAB5549 and BOLD:ACF0272 (Fig. 3B), with a minimum p-distance of 1.08% (~ 7 bp). BOLD:AAB5549 showed some indicators of not being well separated from neighboring BINs with the maximum within-BIN p-distance of 1.77% being higher than the minimum p-distance to the Nearest Neighbor BIN (BOLD:ACF0267) of 1.28%. We performed ASAP clustering and retrieved three different clusters which are also represented in the TCS haplotype network analysis (Fig. 3B). However, we could not find strong morphological traits to separate them, only minor differences between the two BINs (see detailed discussion of those characters under the species description below). And there is no biological data available, as all specimens known from this species have only been collected with Malaise traps. That combined with the molecular differences being rather small (1.28%), does not provide any support to separate the specimens, therefore we here consider the two BINs/ three ASAP clusters to represent the same species.

BIN BOLD:ABZ4155 is shared by two species, D. carlosmanuelrodriguezi (Costa Rica) and D. helenedumasae (Brazil, French Guiana). It has been discussed before (Fernandez-Triana et al. 2019: 100) that this BIN contains more than one species, including some still undescribed from Costa Rica, ACG (Fig. 4A). The full barcode sequence of a paratype of helenedumasae (only specimen with available sequence for this species) is > 1.5% (~ 10 bp) different from the available sequences of carlosmanuelrodriguezi and the remaining (undescribed) Costa Rican species within this BIN. There are also morphological differences between these two species (see details in key below), which we consider to be distinct.

Figure 4.

A TCS haplotype network of BIN BOLD:ABZ4155, sequence length for analysis: 632 bp. Host species are indicated via coloration B TCS haplotype network of BIN BOLD:ABX6267, sequence length for analysis: 651 bp. The different molecular BINs are indicated via coloration.

BIN BOLD:ABX6267 is shared by three Dolichogenidea species, all of them from ACG, richardashleyi, ritaashleyae and yeimycedenoae (Fig. 4B). The differences between these three species ranges between 1.05 and 1.20% (7–8 bp) and they all group in three distinct clusters in the NJ tree. This BIN is also comparatively very close to BOLD:ABY5258, which only contains D. rosamatarritae, another ACG species. The difference between these two BINs is 1.93% (~ 12 bp) compared to a within-BIN maximum p-distance of 1.61% for BOLD:ABX6267 and 0.77% for BOLD:ABY5258. However, there are consistent morphological differences between these four species (Fernandez-Triana et al. 2016, see also key below), and we consider all four to be distinct.

In addition to the cases discussed above, several other BINs may or may not represent a single species and more comprehensive studies would be needed to conclude, ideally including other molecular markers, more morphological analyses, and/or additional biological information. For example, Dolichogenidea rociocordobae and D. frankjoycei, are here considered as distinct because they have been reared from different host species, but the morphological differences are very small and subtle (see couplet 21 in the key below) and DNA barcodes, considered in BOLD as two separate BINs are only 1.01% different (~ 6 bp) (Fig. 5).

Figure 5.

TCS haplotype network of BINs BOLD:ABA3469 and BOLD:ACJ2777, sequence length for analysis: 654 bp. Host species are indicated via coloration.

It could be argued that, in the near future, DNA barcoding data would be the most efficient way to identify species. However, at present DNA barcoding alone is neither sufficient (as one third of the Dolichogenidea here treated do not have available barcodes) nor entirely accurate (as 8.3% of the species with molecular data cannot be unambiguously identified by using BINs). Therefore, morphological and biological data must also be considered to differentiate species.

Parasitoid biology data

Biological data for Neotropical Dolichogenidea are currently incomplete. Only 52 of the species (41.9%) have some sort of host information associated; therefore, the comments below can only be seen as preliminary and may change when biological data are known for more species and more rearings in the region.

There seems to be a prevalence of wasp species with solitary cocoons (76.9%), more than three times the species with gregarious cocoons (23.1%). This is in stark contrast with data for the related genera Apanteles (which had 42% of its Mesoamerican species with solitary cocoons versus 58% gregarious, see Fernandez-Triana et al. 2014a) and Alphomelon (48% of New World species solitary, 52% gregarious, see Fernandez-Triana et al. 2023).

A total of 11 Lepidoptera families were parasitized by Neotropical Dolichogenidea. The most represented group was the microlepidopteran Depressariidae (33.9% of all host data available), followed by Gelechiidae (16.9%), Crambidae (13.6%), Tortricidae (10.2%), Thyrididae (8.5%) and Pyralidae (6.7%). Again, this is in strong contrast with the (morphologically closely related) genus Apanteles, which in Mesoamerica has been recorded from at least 14 families of Lepidoptera but with a completely different arrangement of groups (Fernandez-Triana et al. 2014a): Hesperiidae (33%), Elachistidae (26%), Crambidae (21%), Pyralidae (4%), Choreutidae (3%) and Gelechiidae (3%). The main two groups parasitized by Apanteles (hesperiids and elachistids) are not even found among the known host records of Dolichogenidea in the Neotropics. Although it must be noted that some of the “Gelechiidae” from the Apanteles paper could now be placed in Depressariidae, that still would not make any major difference as that is the family with the smallest percentage of parasitism by Apanteles, whereas it is the largest group attacked by Dolichogenidea. Although more host data would be useful, especially for Dolichogenidea, there seems to be a very distinctive separation between the two parasitoid genera regarding host families used and solitary/gregarious wasp cocoons.

With respect to parasitoid biology, the only similarity between Neotropical species of Dolichogenidea and Apanteles seems to be the fact that both genera are mostly monophagous or at most oligophagous. Out of the 52 Dolichogenidea species with host data available, 29 (55.8%) are only known to parasitize a single host species, whereas 12 (23.1%) parasitize two host species and five (9.6%) parasitize three hosts. Another five wasp species (9.6%) attack either four or five different hosts. Only a single species, D. tomdaleyi, has been recorded parasitizing up to 19 different hosts, but even there that list includes 14 putative species of Antaeotricha and four putative species of Stenoma (both moth genera belonging to the family Depressariidae). In almost all cases, the hosts species belong to one genus (or related genera) in the same Lepidoptera family. Only six Dolichogenidea have host records that include two Lepidoptera families and D. rosamatarritae hosts include three different families. One caveat is that most of the biological information discussed above is based on the extensive rearing data available from ACG material, while data from other Neotropical areas are neither complete nor comprehensive.

Preliminary data on elevational and ecosystem distribution of species

Neotropical Dolichogenidea are here recorded from sea level up to 4,100 m. Although the data available at present are insufficient to draw strong conclusions, the distribution of species appears to be elevationally segregated. Most of the species dealt with in this paper are found between 400–1,500 m; however, a few species have only been found at higher elevations, including a few examples found at more than 3,000 m (Costa Rica) and 4,000–4,100 m in the Andes (South America). Among the species recorded from ACG (the only area where more comprehensive data is available), most species seem to be restricted to one specific ecosystem (dry forest, rain forest or cloud forest).

Key to Dolichogenidea from the Neotropical region

After morphological characters, basic data on species distribution (countries), and, if available or known, biology (solitary/gregarious parasitoid and general info on host), and DNA barcodes (BIN data) are included. The additional data are presented between brackets, and are meant to support and complement (but not supplant) the morphological identification of species. More comprehensive details on species distribution, biology and molecular data are discussed under each species treatment.

1	T2 shape ranging from almost quadrate to broadly rectangular, its width at posterior margin 1.3–2.2× its length centrally, rarely 2.3–2.4×; *and* T2 smooth; *and* ovipositor sheath 1.8–2.5× as long as metatibia length (carlosmanuelrodriguezi group)	2
–	T2 shape ranging from transverse to (rarely) broadly rectangular; its width at posterior margin usually 3.0–5.0× its length centrally, if rarely 2.4–2.9×, *then* T2 strongly sculptured with longitudinal striae or ovipositor sheaths < 1.2× metatibia length; ovipositor sheath usually 0.5–1.5× as long as metatibia length, if very rarely up to 2.0×, *then* T2 strongly sculptured	10
2(1)	Comparatively larger size, body length 4.80–4.90 mm, fore wing length 4.70–4.80 mm, ovipositor sheath length 3.00–3.40 mm; *and* metatrochanter and metatrochantellus entirely black [Costa Rica. Solitary, host: Cerconota recurvella (Depressariidae). BIN BOLD:AAB5701]	D. melaniamunozae Fernandez-Triana & Boudreault, 2019
–	Comparatively smaller size, body length < 4.00 mm (rarely 4.20 mm), fore wing length < 3.70 mm (rarely 4.20 mm), ovipositor sheath length, 3.10 mm (rarely 3.20 mm); *and* metatrochanter and/or metatrochantellus at least partially yellow	3
3(2)	Mesofemur entirely yellow, metafemur almost entirely yellow (except for small, dark spot on anterior 0.1–0.2)	4
–	Mesofemur partially dark brown (rarely mostly yellow with small, dark spots), metafemur mostly to entirely dark brown to black	6
4(3)	Comparatively larger size, body length 4.10–4.20 mm, fore wing length 4.20–4.30 mm, ovipositor sheath length 3.00–3.20 mm [Brazil, French Guiana. BIN BOLD:ABZ4155]	D. helenedumasae Fernandez-Triana & Boudreault, sp. nov.
–	Comparatively smaller size, body length 3.70–3.90 mm, fore wing length 3.70 mm, ovipositor sheath length 2.60–2.90 mm [Costa Rica]	5
5(4)	Slightly smaller specimens (body length 3.70 mm; ovipositor sheath 2.60 mm); T2 posterior width 2.0 × T2 length medially; F2 1.6× as long as F14 [21 DNA barcode diagnostic characters: 31T, 37A, 55G, 67A, 70A, 79A, 88A, 109A, 139A, 190C, 277A, 290G, 316G, 322G, 370T, 386T, 418T, 436T, 460A, 496G, 556A] [Costa Rica. Solitary, host: Antaeotricha spurca (Depressariidae). BIN BOLD:ABZ4155]	D. carlosmanuelrodriguezi Fernandez-Triana & Boudreault, 2019
–	Slightly larger specimens (body length 3.90 mm; ovipositor sheath 2.90 mm); T2 posterior width 1.8 × T2 length medially; F2 1.7–1.9× as long as F14 [21 DNA barcode diagnostic characters: 31A, 37G, 55A, 67T, 70G, 79T, 88T, 109G, 139T, 190A, 277T, 290A, 316A, 322T, 370A, 386A, 418A, 436A, 460G, 496A, 556GA] [Costa Rica. Solitary, host: Stenoma Janzen99 (Depressariidae). BIN BOLD:ABA9255]	D. josealfredohernandezi Fernandez-Triana & Boudreault, 2019
6(3)	T1 comparatively broader, T1 length 1.5 × T1 anterior width, and T1 length 1.5 × T1 posterior width; T2 comparatively less quadrate (T2 posterior width 2.3 × T2 length medially) [Costa Rica. Solitary, host: Brenthia Janzen12 (Choreutidae). BIN BOLD:AAL2298]	D. angelagonzalezae Fernandez-Triana & Boudreault, 2019
–	T1 comparatively narrower, T1 length 1.9–2.4× (very rarely 1.6×) T1 anterior width, and T1 length 1.9–3.0× (very rarely 1.6–1.7× in small specimens) T1 posterior width; T2 comparatively more quadrate, T2 posterior width 1.3–1.9× (very rarely 2.0–2.1× in small specimens) T2 length medially	7
7(6)	Mesofemur mostly yellow, at most with small, dark spots (if mesofemur mostly brown, then specimens significantly smaller, with body length 2.50–3.00 mm)	8
–	Mesofemur mostly dark brown (body length ≥ 3.30 mm, usually more)	9
8(7)	F2 2.1–2.5× (average: 2.3×) as long as F14; T2 posterior width 1.3–1.9× (average: 1.6×) T2 length medially; pterostigma length/width 3.0–3.5× (average 3.2×) [Costa Rica. Gregarious, host: Antaeotricha renselariana (Depressariidae). BIN BOLD:ABX6174]	D. alejandromasisi Fernandez-Triana & Boudreault, 2019
–	F2 1.8–2.1× (average: 1.9×) as long as F14; T2 posterior width 1.6–2.1× (average: 1.8×) T2 length medially; pterostigma length/width 2.7–3.0× (average 2.8×) [Costa Rica, Ecuador. Gregarious, host: Antaeotricha Janzen107 (Depressariidae). BIN BOLD:AAL2325]	D. rogerblancoi Fernandez-Triana & Boudreault, 2019
9(7)	Pterostigma length/width 3.0×; posterior ocellar line 1.6× lateral ocellus diameter; ocular ocellar line 1.2× posterior ocellar line [Costa Rica. Gregarious, host: Antaeotricha phaeoneura (Depressariidae). BIN BOLD:ACC1300]	D. genuarnunezi Fernandez-Triana & Boudreault, 2019
–	Pterostigma length/width 3.1–3.3× (average 3.2×); posterior ocellar line 1.8–2.0× lateral ocellus diameter; ocular ocellar line 1.2–1.5× posterior ocellar line [Costa Rica. Gregarious, host: Antaeotricha Janzen126 (Depressariidae). BIN BOLD:ABY3724]	D. cedenoae Fernandez-Triana & Boudreault, 2025
10(1)	T1 and T2 heavily sculptured with strong longitudinal striae (rarely strong reticulated sculpture) covering entire surface of T2 and most of T1 (at least posterior 0.6–0.9, although centrally there may be a narrow smoother area); T2 often with strong, crenulated sulcus along posterior margin; T2 broadly rectangular or trapezoidal in shape, its width at posterior margin 2.4–2.8× (rarely ~ 3.0×) its length medially or T2 more or less transverse but with anterior and/or posterior margins centrally arcuate (thus increasing T2 length medially, so that width at posterior margin is < or ~ 3.0× its length medially); T1 usually slightly broadening towards posterior margin or parallel-sided, if very rarely T1 mostly parallel-sided but slightly narrowing towards posterior margin near apex *then* T2 always broadly rectangular or trapezoidal	11
–	T1 and T2 much less sculptured, if sculptured rarely with strong longitudinal striae and then only covering margins of posterior half or less on T1 and almost never entire surface of T2 (if very rarely T2 more or less almost entirely sculptured, *then* T2 always very transverse, its width at posterior margin > 3.5× its length medially); T2 rarely with crenulated sulcus along posterior margin (if sulcus present, less strongly impressed and much narrower than above); T2 shape variable but usually transverse (its width at posterior margin > 3.0× its length medially, usually 3.5–4.0× or more); if rarely T2 broadly rectangular or trapezoidal *then* T1 comparatively thin and narrowing towards posterior margin	39
11(10)	T1 comparatively broader, evenly broadening towards posterior margin or mostly parallel-sided but slightly widening towards posterior margin thus T1 length < 1.5× its width at posterior margin; T2 usually broadly rectangular and large, covering most surface of tergum	12
–	T1 comparatively thinner, mostly parallel-sided but posterior 0.1–0.3 slightly narrowing towards posterior margin thus T1 length ≥ 2.0× (rarely 1.7×) its width at posterior margin; T2 usually trapezoidal and rather small, not covering entire surface of tergum (rarely T2 more or less transverse)	30
12(11)	Anteromesoscutum entirely to mostly covered by coarse and deep punctures, face, anterior half of mesopleuron, scutellar disc with coarse punctures; F1–F4 (sometimes F1–F6) yellow to yellow brown, clearly much paler colored than dark brown scape and rest of flagellomeres; T1 and T2 usually black and T4+ usually brown, but metasoma dorsally with some paler coloration always including T3 (which varies from dark brown centrally with orange-yellow laterally, to mostly or entirely yellow or orange-yellow), sometimes also T4 and T5 orange-yellow and rarely even T2 reddish brown; legs mostly yellow or orange-yellow, except for metafemur posterior 0.3–0.5, metatibia 0.3 and entire metatarsus brown to dark brown [Canada, Costa Rica, United States. Solitary (?), hosts: Choristoneura freemani, Epiblema strenuana (Tortricidae), Fissicrambus mutabilis, Neodactria zeellus (Crambidae) BIN BOLD:AAA3764]	D. ensiger (Say, 1836)
–	Anteromesoscutum, face, mesopleuron and scutellar disc with variable sculpture but never as coarse and deep as above; flagellomeres of approx. same coloration and usually brown to dark brown or black; metasoma usually entirely dark brown to black, without extensive pale coloration; legs with more extensive dark coloration	13
13(12)	Ovipositor sheath spatula-shaped and 0.8× as long as metatibia length; distinctive leg color with coxae dark brown to black and rest of legs mostly dark brown (except for bright yellow trochanters and trochantellus, dorsal margin of profemur, anterior 0.1 of pro- and mesotibiae and anterior 0.3 of metatibia; and all tibial spurs white-yellow); tegula dark brown to black, humeral complex mostly dark brown; pterostigma bright yellow-white but with thin brown margins, most of wing veins pale yellow-brown; F15 sub-cubic (1.1× as long as high); propodeum areola mostly defined, but open on anterior ~ 0.3, and without defined transverse carinae; body length 3.80 mm, fore wing length 4.20 mm [Colombia]	D. caldas Fernandez-Triana & Boudreault, sp. nov.
–	Ovipositor sheath not spatula-shaped; different leg coloration, usually neither as dark nor as contrasting; tegula and humeral complex variable but not as dark as above; pterostigma and wing venation color not as above; F15 usually not cubic, often > 1.3× as long as high; propodeum areola variable but not as above; body and fore wing lengths variable but usually much smaller	14
14(13)	Metatibia dark brown to black on posterior 0.5–0.8; body length comparatively smaller, usually 2.20–2.70 mm	15
–	Metatibia mostly yellow with only dark brown to black on posterior 0.1–0.2 (in addition there can be a dark brown band dorsally on entire length of metatibia); body length comparatively larger, usually 3.10–3.80 mm	24
15(14)	Metatibia dark brown to black on posterior 0.5; ocular ocellar line 1.4× posterior ocellar line	16
–	Metatibia dark brown to black on posterior 0.6–0.8; ocular ocellar line 1.0–1.1× posterior ocellar line	17
16(15)	F15 length/width 1.5×; size comparatively smaller, body and fore wing lengths 2.30–2.40 mm; pterostigma brown [Costa Rica, 1,000 m]	D. monocavus (Valerio & Whitfield, 2004)
–	F15 length/width 1.1×; size comparatively larger, body length 2.70 mm, fore wing length 3.00 mm; pterostigma brown with small pale spot at base [Venezuela, 1,900 m]	D. verobrondexae Fernandez-Triana & Boudreault, sp. nov.
17(15)	T3 sculptured on anterior 0.2–0.5 (but there mostly centrally); T1 and T2 mostly with reticulate sculpture; tarsal claws of hind legs simple [Costa Rica. Solitary, hosts: Brenthia Janzen05, Stenoma Phillips543 (Choreutidae, Gelechiidae, Depressariidae). BIN BOLD:ABY5258]	D. rosamatarritae (Fernandez-Triana, 2016)
–	T3 entirely smooth; T1 and T2 mostly with longitudinally striated sculpture; tarsal claws of hind legs with single spine-like basally	18
18(17)	Metafemur yellow on anterior half and brown on posterior half; and ovipositor sinuate; and all laterotergites, sternites and hypopygium yellow; and T3–T6 centrally brown, laterally with yellow spots [Ecuador]	D. palenque Fernandez-Triana & Boudreault, sp. nov.
–	Metafemur color different, either mostly brown or entirely to almost entirely yellow; and ovipositor not sinuate; metasoma color variable but not as above	19
19(18)	Metafemur mostly brown (only anterior 0.1 and posterior 0.1 yellow)	20
–	Metafemur entirely yellow, at most with brown spot dorsally on posterior 0.2 or less	22
20(19)	F15 2.0× as long as wide; DNA barcodes 5.9% different from closest BIN and > 7.0% different from species in second half of couplet [Costa Rica. BIN BOLD:AAM5843]	D. rodrigogamezi Fernandez-Triana & Boudreault, sp. nov.
–	F15 1.5× as long as wide; DNA barcodes > 7.0% different from species in first half of couplet [BIN BOLD:ACJ2777, BIN BOLD:ABA3469]	21
21(20)	Ovipositor sheath almost as long as metatibia length (0.95×); F15 comparatively shorter, 1.2× as long as wide [Costa Rica. Solitary, hosts: Dichomeris spp. (Gelechiidae). BIN BOLD:ACJ2777]	D. rociocordobae Fernandez-Triana & Boudreault, sp. nov.
–	Ovipositor sheath clearly shorter than metatibia length (0.80×); F15 comparatively longer, 1.4× as long as wide [Costa Rica. Solitary, hosts: Platynota spp. (Tortricidae). BIN BOLD:ABA3469]	D. frankjoycei Fernandez-Triana & Boudreault, sp. nov.
22(19)	Tegula and humeral complex of same color, white-yellow [Costa Rica. Solitary, host: undetermined species of Gelechiidae with interim name ‘gelJanzen01 Janzen319’. BIN BOLD:AAN2496]	D. mariabustosae (Fernandez-Triana, 2016)
–	Tegula white-yellow, clearly paler than brown humeral complex	23
23(22)	Pro- and mesocoxae yellow; laterotergites 1–4 and at least sternites 1 and 2 entirely to mostly yellow; pterostigma with very small pale spot on anterior 0.1 or less; F15 length 1.4–1.5× its height [Costa Rica, 600–700 m. Solitary, host: Gonionota Janzen116 (Depressariidae). BIN BOLD:ACC4119]	D. felipechavarriai Fernandez-Triana & Boudreault, sp. nov.
–	Pro- and mesocoxae dark reddish brown; all laterotergites and sternites pale brown to dark brown; pterostigma with pale spot on anterior 0.25; F15 length 1.2× its height [Guatemala, 2,000 m]	D. chichicastenango Fernandez-Triana & Boudreault, sp. nov.
24(14)	Pterostigma with pale yellow spot on proximal 0.5; smaller species (body length 3.10 mm and fore wing length 3.40 mm) [Costa Rica. Solitary, host: undetermined species of Gelechiidae with interim name ‘gelJanzen01 Janzen349’. BIN BOLD:ABX6267]	D. richardashleyi (Fernandez-Triana, 2016)
–	Pterostigma mostly brown, at most with small pale spot on proximal 0.1–0.2; usually, but not always, larger species (body length 3.30–3.80 mm and fore wing length 3.50–4.10 mm)	25
25(24)	Metatibial spurs with brown tips; F2 length 2.1 × F14 length [Costa Rica. BIN BOLD:ABX6267]	D. yeimycedenoae (Fernandez-Triana, 2016)
–	Metatibial spurs entirely yellow; F2 length 2.5 × F14 length	26
26(25)	T2 more or less transverse, with anterior margin centrally arcuate, so that its width at posterior margin is ~ 3.0× its length medially; and metacoxa almost entirely dark brown (very small yellow spot on posterior 0.1); and body and fore wing lengths 3.50–3.70 mm [Colombia]	D. antioquia Fernandez-Triana & Boudreault, sp. nov.
–	T2 broader, its width at posterior margin ≤ 2.8× its length medially (usually less); *either* metacoxa dark brown on anterior half and yellow on posterior half, or metacoxa almost entirely dark brown *and* body and fore wing lengths ≤ 3.30 mm	27
27(26)	Hind legs tarsal claws simple; metacoxa dark brown on anterior half and yellow on posterior half; comparatively larger species, body length 3.50–3.80 mm	28
–	Hind legs tarsal claws with single spine; metacoxa almost entirely dark brown (very small yellow spot on posterior 0.1); comparatively smaller species, body length ≤ 3.30 mm	29
28(27)	Metatibia with dorsal dark brown band on entire length of metatibia; pterostigma mostly pale yellow-brown but with thin brown margins; T1 comparatively narrower, ~ 2.0× as long as wide at posterior margin [Costa Rica]	D. heredia Fernandez-Triana & Boudreault, sp. nov.
–	Metatibia without dorsal dark band; pterostigma mostly brown, with pale spot at base; T1 comparatively broader, < 1.6× as long as wide at posterior margin [Costa Rica. BIN BOLD:ABX6267]	D. ritaashleyae (Fernandez-Triana, 2016)
29(27)	Metatibia with dark brown spot on posterior 0.2; metatarsus mostly brown; T2 more transverse, its width at posterior margin > 3.0× its central length; comparatively smaller species, body length 2.70–2.90 mm; ovipositor sheath 1.0–1.1× metatibia length [Costa Rica. Solitary, hosts: Depressariidae: elachJanzen01 Janzen409; Gelechiidae: Dichomeris Janzen76, gelJanzen01 Janzen116. BIN BOLD:AAC5949]	D. tiboshartae Fernandez-Triana & Boudreault, sp. nov.
–	Metatibia almost entirely yellow, with pale brown spot on posterior 0.1; metatarsus mostly yellow to yellow-pale brown; T2 subrectangular its width at posterior margin < 2.7× its central length; comparatively larger species, body length 3.10–3.30 mm; ovipositor sheath 1.2–1.3× metatibia length [Costa Rica. Solitary, hosts: Megalota spp. (Tortricidae). BIN BOLD:AAY4690]	D. ninamasisae Fernandez-Triana & Boudreault, sp. nov.
30(11)	Ovipositor sheath 1.6× metatibia length; and T1 length 1.8× its width at posterior margin; and legs almost entirely brown to dark brown (except for yellow protibial and protarsus and very small, paler spots on posterior 0.1 of pro- and mesofemora and anterior 0.1–0.2 of meso- and metatibiae); and tegula and humeral complex dark brown; and pterostigma mostly yellow-white but with thin brown margins [Guatemala]	D. alexandrei Fernandez-Triana & Boudreault, sp. nov.
–	Ovipositor sheath much shorter or much longer than above; and/or T1 length almost always > 2.0× its width at posterior margin; and/or legs with at least some segments pale colored (yellow to yellow-white or yellow-orange); and/or tegula and/or humeral complex usually yellow; and/or pterostigma color variable but usually not as above	31
31(30)	Ovipositor sheath ~ 2.0× metatibia length; body length 3.20–3.50 mm	32
–	Ovipositor sheath ≤ 1.2× metatibia length (usually much less); usually body length 2.30–2.40 mm (rarely 2.70–3.20 mm)	33
32(31)	Hypopygium and most sternites dark brown; metacoxa partially yellow partially dark brown; comparatively larger ovipositor sheath (1.70–2.00 mm) in spite of slightly smaller body and fore wing (2.60–3.10 mm) than below, with ratio of ovipositor sheath length/body length of 0.60–0.65 [Costa Rica. Solitary, hosts: Antaeotricha spp. Chlamydastis Janzen04, Stenoma spp. (Depressariidae and Crambidae). BIN BOLD:AAD8952]	D. tomdaleyi Fernandez-Triana & Boudreault, sp. nov.
–	Hypopygium and all sternites yellow; metacoxa entirely brown; comparatively shorter ovipositor sheath (1.50 mm) despite slightly larger sized body and fore wing (3.20 mm) than above, with ratio of ovipositor sheath length/body length of 0.47 [Costa Rica. Solitary, host: undetermined species of Depressariidae with interim name elachBioLep01 BioLep286]	D. sallydaleyae Fernandez-Triana & Boudreault, sp. nov.
33(31)	All legs almost entirely brown to dark brown (except for very small, paler spots, on posterior 0.1 of pro- and mesofemora and anterior 0.1–0.2 of tibiae); wings slightly infumated; tegula and humeral complex brown; F15 cubic (around same length than width); T2 shape trapezoidal but rather narrow, barely wider than T1; pterostigma mostly yellow-white but with thin brown margins [Bolivia]	D. yungas Fernandez-Triana & Boudreault, sp. nov.
–	Legs with at least some segments pale colored (yellow to yellow-white or yellow-orange); wings not infumated; tegula and/or humeral complex usually yellow; F15 not cubic (longer than wide); if T2 trapezoidal not as narrow as above; pterostigma color variable but usually not as above	34
34(33)	Tegula brown; ovipositor strongly sinuate; propodeum areola narrower (its height > 1.5× its central width) and open anteriorly [Costa Rica. BIN BOLD:AAM5848]	D. carlosalvaradoi Fernandez-Triana & Boudreault, sp. nov.
–	Tegula yellow; ovipositor not sinuate; propodeum areola broader (its height ~ 1.2× its central width) and close anteriorly	35
35(34)	Scutellar disc with coarse punctures; anterior half of mesopleuron and anteromesoscutum with relatively coarse punctures; pterostigma pale brown but centrally paler than margins; tegula and humeral complex yellow; metatibia entirely brown to dark brown [Costa Rica. BIN BOLD:AAD2236]	D. claudiadoblesae Fernandez-Triana & Boudreault, sp. nov.
–	Scutellar disc smooth; anterior half of mesopleuron smooth, anteromesoscutum with sparse and relatively shallow punctures; pterostigma mostly pale brown with small, paler spot anteriorly; tegula white-yellow, humeral complex mostly brown; metatibia yellow at least on anterior 0.4	36
36(35)	T3 with yellow spots laterally, centrally pale brown, T4+ pale brown to brown; laterotergites 1–5 yellow, most sternites at least partially yellow, hypopygium partially yellow and partially pale brown; pro- and mesocoxae entirely yellow, metacoxa reddish brown; propodeum with carinae defining areola strongly risen and sharp; F15 1.3× as long as high [Ecuador]	D. isabelleae Fernandez-Triana & Boudreault, sp. nov.
–	Metasoma dorsally entirely dark brown to black; all laterotergites, sternites and hypopygium entirely brown to dark brown; pro- and mesocoxae partially reddish brown to brown, metacoxa black or dark brown; propodeum with carinae defining areola not as strongly risen as above; F15 ≥ 1.5× as long as high [Costa Rica]	37
37(36)	Ovipositor comparatively very thin, much thinner than half flagellomeres width; T2 comparatively less transverse, its width at posterior margin 2.8× its central length; metatibia yellow on anterior 0.4–0.5 [Costa Rica. BIN BOLD:AAM5846]	D. jorgecortesi Fernandez-Triana & Boudreault, sp. nov.
–	Ovipositor comparatively thicker, at least as thick as 0.8× flagellomeres width; T2 comparatively more transverse, its width at posterior margin > 3.1× its central length; metatibia mostly yellow, only posterior 0.2 or less dark brown	38
38(37)	Comparatively smaller species, body length 2.30 mm; parasitizing Erebidae; wasp cocoons solitary; DNA barcodes 3.7% different from closest BIN and 5.3% different from species in second half of couplet [Costa Rica. Solitary, host: Rivula Poole03BIN (Erebidae). BIN BOLD:ABA7252]	D. andreamezae Fernandez-Triana & Boudreault, sp. nov.
–	Comparatively larger species, body length 2.70–3.00 mm; parasitizing Mimallonidae; wasp cocoons gregarious; DNA barcodes 2.1% different from closest BIN and 5.3% different from species in first half of couplet [Costa Rica. Gregarious, host: Eadmuna Janzen01 (Mimallonidae). BIN BOLD:AAB4946]	D. pedroleoni Fernandez-Triana & Boudreault, sp. nov.
39(10)	Pro- and mesocoxae pale (white, yellow, orange, yellow-pale brown); metacoxa pale entirely or mostly pale (anterior 0.3–0.4 dark), very rarely metacoxa mostly dark brown	40
–	Pro- and mesocoxae mostly to entirely dark (dark orange-brown, brown, dark brown, black); and metacoxa entirely (often) or mostly (rarely) dark, at most with small pale area on 0.2 or less of metacoxa	65
40(39)	Comparatively paler colored species; T3 (sometimes also T4 and part of T5) entirely yellow to yellow-white, rest of tergites pale brown (except for dark brown T1); and hypopygium and all or most laterotergites and sternites yellow to yellow-white; and all coxae yellow-white, rest of legs mostly yellow to yellow-white (except for part of metafemur and metatibia); and antenna pale brown; and tegula and humeral complex yellow; and mesosoma with some spots pale brown to orange-brown; and fore wing veins r and 2RS strongly angulate; and flagellomeres with comparatively long setae, as long as half flagellomere width	41
–	Comparatively darker colored species, usually all or most tergites brown to black, if rarely with T1 or T2 yellow or pale brown-yellow then T3 always brown, or T3 laterally yellow but centrally brown; laterotergites, sternites and hypopygium variable but very rarely all pale; and/or legs usually with some segments dark; antenna color usually dark brown to black, rarely pale brown; tegula and humeral complex color variable; fore wing veins r and 2RS variable but rarely strongly angulate; flagellomeres usually with shorter setae	43
41(40)	T1 more or less parallel-sided, only very slightly narrowing near posterior margin; T4 (mostly) and T5 (partially) yellow to yellow-white; metafemur mostly yellow-white with only posterior 0.1 brown [Ecuador]	D. limoncocha Fernandez-Triana & Boudreault, sp. nov.
–	T1 strongly narrowing towards posterior margin so that width at anterior margin is clearly longer than width at posterior margin; most tergites brown to reddish brown (except for yellow T3); metafemur either entirely yellow or yellow on anterior 0.5 and dark brown on posterior 0.5	42
42(41)	Flagellomeres orange-yellow; metafemur entirely yellow; T1 very strongly narrowing posteriorly, its width at anterior margin ~ 2.0× its width at posterior margin [Peru. Solitary, host: Microthyris anormalis (Crambidae)]	D. croceicornis (Muesebeck, 1958)
–	Flagellomeres brown; metafemur yellow on anterior 0.5 and dark brown on posterior 0.5; T1 not as strongly narrowing posteriorly, its width at anterior margin ~ 1.5× its width at posterior margin [Costa Rica, Ecuador, Venezuela. Solitary, hosts: Herpetogramma Janzen04, Rhectocraspeda Solis05 (Crambidae). BIN BOLD:AAF5364]	D. alexamasisae Fernandez-Triana & Boudreault, sp. nov.
43(40)	T3 mostly yellow (only pale brown on central part); and first and second pair of legs almost entirely yellow (except for pale brown mesocoxa), third pair of legs mostly brown to dark brown (except for yellow trochanter, trochantellus, anterior 0.1 and posterior 0.1 of metafemur, and anterior 0.5–0.6 of metatibia); T1 mostly strongly sculptured, T2 entirely smooth; T2 comparatively narrow and sub-quadrate, its width at posterior margin 2.0× its central length; fore wing veins r and 2RS strongly angulate; pterostigma mostly yellow-white with thin brown margins [Peru, 2,000 m]	D. amazonas Fernandez-Triana & Boudreault, sp. nov.
–	T3 entirely pale brown, dark brown or black; leg coloration different than above; and/or T1 and T2 sculptured different than above; and/or T2 more transverse; and/or fore wing veins r and 2RS variable but rarely strongly angulate; and/or pterostigma coloration different than above	44
44(43)	Posterior 0.1–0.2 of T1 (centrally) and entire T2 yellow to pale brown-yellow; T3+ pale brown; metacoxa entirely to almost entirely dark brown; anteromesoscutum with coarse, deep and dense punctures (separation between punctures less than individual puncture diameter); face, propleuron, pronotum, most of mesopleuron, scutellar disc, and most of outer side of metacoxa mostly to entirely covered by relatively coarse punctures [Costa Rica. BIN BOLD:AAF7717]	D. ericpalolai Fernandez-Triana & Boudreault, sp. nov.
–	Different coloration pattern of T1 and T2 (usually mostly to entirely brown, dark brown or black); T3+ almost always dark brown to black; metacoxa usually entirely to partially pale colored (white, yellow-white, yellow, orange-yellow); anteromesoscutum rarely as strongly sculptured; face, propleuron, pronotum, most of mesopleuron, scutellar disc and most of outer side of metacoxa with variable sculpture, but usually mostly smooth or with shallow and/or sparse punctures	45
45(44)	Metatibia entirely to mostly pale (at most with darker area on posterior 0.3 or less), if metatibia with brown margin dorsally then rest of legs yellow or white-yellow	46
–	Metatibia entirely to mostly dark (at most with paler area on anterior 0.3 or less) *and* at least some other leg segments dark	58
46(45)	Extensive orange coloration, including clypeus, anteromesoscutum (almost entirely, except for small dark spot near scutellar disc), antero-dorsal spot on mesopleuron, propleuron (partially), mesosternum (mostly) and two faint spots postero-laterally on T1 [Costa Rica. BIN BOLD:AAM5736]	D. shelleymcsweeneyae Fernandez-Triana & Boudreault, sp. nov.
–	If there is any orange or orange-yellow coloration, it is very limited to small, isolated spots	47
47(46)	Metafemur and metatibia mostly yellow-white but with brown to dark brown margins dorsally (dark dorsal margin only partially defined on metafermur); pterostigma mostly white-yellow on anterior 0.5–0.7, with darker (pale brown) margins [Costa Rica. Solitary, host: undetermined Crambidae. BIN BOLD:ABY1812]	D. anikenpalolae Fernandez-Triana & Boudreault, sp. nov.
–	Metafemur and metatibia mostly yellow or yellow-white (at most with brown spot on posterior 0.3 or less), without dark margins dorsally; pterostigma uniformly colored, mostly brown to brown, or brown with small pale spot on anterior 0.2	48
48(47)	T1 strongly narrowing towards posterior margin, its length medially ~ 4.0–5.0× its width at posterior margin and its width at anterior margin 2.0× its width at posterior margin	49
–	T1 either parallel-sided or mostly parallel-sided and only narrowing towards posterior margin near its posterior half, its length medially ≤ 3.0× its width at posterior margin (usually less) and its width at anterior margin < 1.5× its width at posterior margin (usually less)	51
49(48)	Humeral complex, all laterotergites, sternites and hypopygium brown to dark brown [Costa Rica. BIN BOLD:AAI9740]	D. tristanpalolai Fernandez-Triana & Boudreault, sp. nov.
–	Humeral complex, at least some laterotergites, all sternites and hypopygium yellow	50
50(49)	Ovipositor sheath almost as long as metatibia length (0.9×); tegula yellow, same color than humeral complex [Costa Rica. BIN BOLD:AAD6850]	D. junhyongkimi Fernandez-Triana & Boudreault, sp. nov.
–	Ovipositor sheath clearly shorter than metatibia length (0.7×); tegula brown, darker than yellow humeral complex [Costa Rica. BIN BOLD:AAM5850]	D. ingredolsonae Fernandez-Triana & Boudreault, sp. nov.
51(48)	Ovipositor comparatively thicker, as wide or wider than flagellomeres width; and F15 comparatively very long, 2.0× as long as its width; and fore wing vein r longer than pterostigma height and ~ 2.0× as long as vein 2RS; and all legs yellow or orange yellow, except for metacoxa mostly dark brown to black but with posterior 0.2 yellow; and pterostigma mostly brown but with pale spot at anterior 0.2 [French Guiana]	D. yvesbraeti Fernandez-Triana & Boudreault, sp. nov.
–	Ovipositor comparatively thinner, its width much less than flagellomeres width; and/or F15 comparatively shorter, < 1.5× as long as its width; and/or fore wing vein r not approx. same length or shorter than pterostigma height and < 1.5× as long as vein 2RS; and/or legs with different coloration pattern; and pterostigma mostly brown to pale brown	52
52(51)	T2 comparatively less transverse, its length 3.0× its width at posterior margin	53
–	T2 comparatively more transverse, its length 4.0–5.0× its width at posterior margin	54
53(52)	Ovipositor sheath 1.0× as long as metatibia; fore wing veins r and 2RS not meeting at strong angle; comparatively smaller species, body length 2.00 mm, fore wing length 2.20 mm [Costa Rica, Honduras. BIN BOLD:ACE8228]	D. katiemccluskeyae Fernandez-Triana & Boudreault, sp. nov.
–	Ovipositor sheath 0.75× as long as metatibia; fore wing veins r and 2RS meeting at a strong angle; comparatively larger species, body length 2.60 mm, fore wing length 2.70 mm [Costa Rica]	D. arenal Fernandez-Triana & Boudreault, sp. nov.
54(52)	Tegula and humeral complex dark brown [Costa Rica. BIN BOLD:AAJ1396]	D. leahdennisae Fernandez-Triana & Boudreault, sp. nov.
–	Tegula yellow or pale brown, humeral complex yellow	55
55(54)	Scutellar disc mostly with punctures; anteromesoscutum with relatively coarse punctures; T1 length medially ~ 3.0× its width at posterior margin; anterior 0.3 of mesopleuron and posterior 0.4–0.5 of metapleuron sculptured [Costa Rica. BIN BOLD:AAC7481]	D. joshdarfleri Fernandez-Triana & Boudreault, sp. nov.
–	Scutellar disc usually smooth and shiny, without punctures (at most with very shallow punctures along margins); anteromesoscutum mostly smooth or with relatively shallow punctures; T1 length medially ≤ 2.5× its width at posterior margin (usually less); mesopleuron and metapleuron entirely to almost entirely smooth or with few, shallow punctures	56
56(55)	T2 mostly smooth; T1 often (but not always) with anterior 0.4–0.5 yellow-brown, clearly paler colored than rest of tergite (which is dark brown or reddish brown) [Costa Rica. Solitary, host: Antaeotricha Janzen221 (Depressariidae). BIN BOLD:ABX5195]	D. robinsherwoodae Fernandez-Triana & Boudreault, sp. nov.
–	T2 entirely sculptured (rarely weakly sculptured but then sculpture covering most of tergite); T1 entirely dark brown or black	57
57(56)	Scape yellow, much paler than brown flagellomeres; ovipositor sheath 0.7× as long as metatibia; pterostigma entirely pale brown [Canada, Honduras, United States. Solitary, hosts: Keiferia spp. (Gelechiidae)]	D. phthorimaeae (Muesebeck, 1921)
–	Scape brown, same color than flagellomeres; ovipositor sheath 0.9× as long as metatibia; pterostigma mostly bright yellow-white (but with central, darker spot which is pale brown) [Argentina]	D. tucuman Fernandez-Triana & Boudreault, sp. nov.
58(45)	T1 comparatively narrow, its length medially ~ 3.0× its width at posterior margin; and tegula brown, clearly darker than yellow humeral complex; and all trochanters, pro- and mesocoxae white; and metacoxa mostly white-yellow, with dark brown spot on anterior 0.2–0.3 [Costa Rica. BIN BOLD:ACF2929]	D. paulfryi Fernandez-Triana & Boudreault, sp. nov.
–	T1 usually comparatively broader, its length medially usually 2.0–2.5× its width at posterior margin (rarely up to 3.0×); and/or tegula white or yellow, similar color or paler than humeral complex; and/or all trochanters, pro- and mesocoxae yellow; and metacoxa entirely to mostly dark brown (very rarely mostly yellow with dark brown spot on anterior 0.3)	59
59(58)	Metafemur entirely to mostly yellow (at most thin brown area dorsally on posterior 0.3 or less); T2 with large, smooth area centrally, sculpture limited to margins	60
–	Metafemur mostly dark brown (except for yellow-white spot on anterior 0.2–0.3 or posterior 0.2); T2 mostly sculptured, at most with small central area smooth	61
60(59)	T1 comparatively narrower, its length medially ~ 3.0× its width at posterior margin [Costa Rica. BINs BOLD:AAB5549, BOLD:ACF0272]	D. annlisterudae Fernandez-Triana & Boudreault, sp. nov.
–	T1 comparatively broader, its length medially ~ 2.2× its width at posterior margin [Costa Rica. BIN BOLD:ACF0267]	D. johnrobinsoni Fernandez-Triana & Boudreault, sp. nov.
61(59)	Pterostigma entirely yellow-white or mostly yellow-white but with thin brown margins; metacoxa entirely dark brown	62
–	Pterostigma brown; metacoxa half brown and half pale (yellow or yellow-white)	63
62(61)	Much larger body size (body length 4.00 mm, fore wing length 4.50 mm); all sternites yellow, hypopygium yellow on anterior 0.5, dark brown on posterior 0.5; pterostigma mostly yellow-white but with thin brown margins; T1 slightly broadening towards posterior margin, its length 1.2× its width at posterior margin; T1 almost entirely covered by coriaceous sculpture; T2 comparatively less transverse, its width at posterior margin 3.0× its central length; T2 mostly covered by longitudinal sculpture, but with two smoother, small areas near anterior margin centrally [Ecuador]	D. papallacta Fernandez-Triana & Boudreault, sp. nov.
–	Much smaller body size (body length and fore wing length 2.20–2.50 mm); all sternites and hypopygium brown; pterostigma entirely yellow-white; T1 distinctively narrowing from posterior 0.3 towards posterior margin, its length 2.0× its width at posterior margin; T2 comparatively more transverse, its width at posterior margin > 3.5× its central length, its length 2.0× its width at posterior margin; T1 posterior 0.6 and T2 (mostly) sculptured [Costa Rica. BIN BOLD:AAM5842]	D. teremariae Fernandez-Triana & Boudreault, sp. nov.
63(61)	T2 with central area smooth; tegula yellow, same color than humeral complex; most of metafemur and metatibia brown [Costa Rica. BIN BOLD:AAM5088]	D. jennyphillipsae Fernandez-Triana & Boudreault, sp. nov.
–	T2 mostly sculptured (if there is a central area smooth, it is very small); tegula white-yellow, clearly paler in color than yellow humeral complex; most of metafemur and metatibia black	64
64(63)	DNA barcodes with 14 diagnostic characters: 100G, 115C, 127A, 197T, 202T, 313A, 316G, 322A, 340C, 343C, 388G, 397C, 457C, 484C [Costa Rica. BIN BOLD:AAC8392]	D. robertofernandezi Fernandez-Triana & Boudreault, sp. nov.
–	DNA barcodes with 14 diagnostic characters: 100A, 115T, 127T, 197C, 202C, 313G, 316A, 322G, 340T, 343T, 388A, 397T, 457T, 484T [Costa Rica. BIN BOLD:AAB9372]	D. isidrochaconi Fernandez-Triana & Boudreault, sp. nov.
65(39)	Metafemur brown on anterior half and yellow on posterior half; and ovipositor sheath 0.6× as long as metatibia length; and propodeum mostly smooth with only two carinae partially defining an areola on posterior 0.4; and T1 mostly sculptured on posterior 0.6 and T2 mostly smooth [Chile]	D. ericsimoni Fernandez-Triana & Boudreault, sp. nov.
–	Metafemur mostly pale or mostly dark; ovipositor sheath variable but often longer than 0.6× metatibia length; propodeum variable but often with some sculpture in addition to propodeal carinae; sculpture of T1 and T2 variable but often not as above	66
66(65)	Metafemur entirely to mostly pale (at most with darker spot on posterior 0.3 or less, or narrow dark borders around margins but still mostly pale)	67
–	Metafemur entirely to mostly dark (at most with pale spots on anterior 0.2 or less, or posterior 0.2 or less, or with very small area centrally paler colored)	78
67(66)	Ovipositor sheath < 0.5× metatibia length	68
–	Ovipositor sheath > 0.7× metatibia length (usually much more)	69
68(67)	DNA barcodes with 64 diagnostic characters: 56T, 67T, 73A, 88A, 127A, 133T, 139A, 172A, 181G, 184A, 190T, 205A, 211G, 217A, 220A, 223T, 229A, 232A, 241G, 244G, 256A, 274T, 278T, 281T, 283T, 293T, 298T, 311A, 312C, 319G, 322G, 325A, 343A, 352T, 356A, 357T, 358T, 359T, 368G, 376A, 386T, 400T, 412T, 421A, 433T, 434G, 436T, 439T, 445T, 448T, 449A, 454A, 455A, 482A, 484A, 494A, 526T, 528C, 529A, 544G, 547T, 553T, 568T, 574T [Costa Rica. BIN BOLD:AAE8596]	D. bernardoespinozai Fernandez-Triana & Boudreault, sp. nov.
–	DNA barcodes with 64 diagnostic characters: 56A, 67A, 73G, 88T, 127T, 133C, 139T, 172T, 181A, 184G, 190A, 205T, 211A, 217T, 220T, 223A, 229G, 232T, 241A, 244A, 256T, 274A, 278G, 281G, 283A, 293A, 298A, 311G, 312T, 319A, 322T, 325T, 343T, 352A, 356T, 357C, 358G, 359A, 368A, 376G, 386A, 400A, 412A, 421T, 433G, 434T, 436A, 439A, 445C, 448A, 449T, 454T, 455G, 482T, 484G, 494T, 526A, 528T, 529T, 544A, 547A, 553A, 568A, 574A [Costa Rica. Solitary, host: Immidae. BIN BOLD:ACI3413]	D. angelsolisi Fernandez-Triana & Boudreault, sp. nov.
69(67)	Posterior 0.5 of T1 and entire T2 sculptured with strong, longitudinal striae; and T1 comparatively thinner, its length 2.0× its width at posterior margin; and T2 width at posterior margin 3.7× its length medially; and body color mostly pale reddish brown; and pterostigma with small pale (pale yellow) with white spots on anterior 0.1 and posterior 0.1; and propodeum mostly smooth, with comparatively tall and thin areola that occupies the entire length of propodeum and it is completely defined by carinae; and ovipositor sheath 1.5× metatibia length [Colombia]	D. putumayo Fernandez-Triana & Boudreault, sp. nov.
–	Different combination of characters, either with posterior 0.5–0.6 of T1 and/or T2 mostly smooth or T2 sculptured but with small polished area centrally; and/or T2 width at posterior margin < 3.0× its length medially; and body color mostly dark brown to black; and pterostigma with different color pattern; and/or propodeum with areola variously defined and not as thin; and/or ovipositor sheath length either shorter or longer than 1.5× metatibia length	70
70(69)	T1 strongly narrowing towards posterior margin (width at anterior margin 1.8× width at posterior margin), T1 length 4.5× its width at posterior margin; and T2 weakly sculptured and trapezoidal, its width at posterior margin 2.5× its central length [Brazil]	D. escobarae Fernandez-Triana & Boudreault, sp. nov.
–	T1 broader, its width at anterior margin similar or only very slightly broader (1.2× or less) than width at posterior margin, T1 length < 3.0× its width at posterior margin; T2 sculpture and shape variable but not as above	71
71(70)	Posterior 0.5–0.6 of T1 and/or T2 mostly smooth, if sculptured, without strong longitudinal striae mostly covering terga surface; and/or ovipositor sheath < 1.2× as metatibia length; and/or T2 transverse and comparatively narrow, its width at posterior margin > 3.0× its length medially (usually more)	72
–	Posterior 0.5–0.6 of T1 and T2 mostly with strong sculpture, usually longitudinal striae mostly covering terga surface (but T2 with small polished area centrally); and ovipositor comparatively very long, ≥ 1.5× as metatibia length (usually much more, rarely 1.3–1.4×); T2 more or less transverse but with anterior and/or posterior margins strongly arcuate, so that T2 length is longer medially than laterally and thus T2 width at posterior margin is usually < 3.0× its length medially	75
72(71)	Propodeum without areola, with only small carinae near nucha; metatarsus yellow-brown, similar color than metatibia [Saint Vincent]	D. parallelis (Ashmead, 1900)
–	Propodeum with areola, areola delimited by carina posteriorly and laterally; metatarsus dark brown to black, clearly much darker than yellow-brown metatibia [Costa Rica]	73
73(72)	T1 shiny and almost entirely smooth (at most with few shallow punctures along lateral margins on posterior 0.3); T1 broader, its length medially 2.0× its width at posterior margin [Costa Rica. Solitary, hosts: Chlamydastis spp., Stenoma Janzen199 (Depressariidae). BIN BOLD:AAT8860]	D. ronaldzunigai Fernandez-Triana & Boudreault, sp. nov.
–	T1 with some sculpture on posterior half; T1 narrower, length medially ≥ 2.5× its width at posterior margin	74
74(73)	Scape entirely dark brown to black; pro- and mesocoxae brown, metacoxa black; tegula yellow, much paler than brown humeral complex; anteromesoscutum punctures near end of notauli well separated and similar to puntures on rest of anteromesoscutum [Costa Rica. BIN BOLD:AAM5740]	D. juanmatai Fernandez-Triana & Boudreault, sp. nov.
–	Scape ventrally yellow-brown, distinctly paler colored than dorsal side; pro- and mesocoxae partially pale brown partially yellow, metacoxa mostly dark brown to black but with posterior 0.1–0.2 yellow; tegula brown, same color than humeral complex; anteromesoscutum punctures near end of notauli fused, unlike punctures on rest of anteromesoscutum [Costa Rica. Gregarious, hosts: Herpetogramma spp. (Crambidae)]	D. jaimelewisi Fernandez-Triana & Boudreault, sp. nov.
75(71)	Vannal lobe straight and with very few, very small setae [United States. Solitary, hosts: Acrobasis caryae, Gretchena bolliana (Pyralidae, Tortricidae)]	D. acrobasidis (Muesebeck), 1921
–	Vannal lobe slightly convex and uniformly bordered by setae	76
76(75)	Ovipositor very long, 2.0× as long as metatibia length; pterostigma with relatively large pale (yellow-white) spot at base that occupies 0.3–0.4 pterostigma length [Costa Rica. Solitary, host: Ategumia lotanalis (Crambidae). BIN BOLD:ABY7999]	D. anniapicadoae Fernandez-Triana & Boudreault, sp. nov.
–	Ovipositor much shorter, ≤ 1.4× as long as metatibia length; pterostigma usually without pale spot at base or with small pale spot occupying < 0.1 pterostigma length	77
77(76)	Tegula and humeral complex yellow; most laterotergites, some sternites and sometimes hypopygium yellow to yellow-brown [Costa Rica. BIN BOLD:AAM5847]	D. jorgecarvajali Fernandez-Triana & Boudreault, sp. nov.
–	Tegula dark brown, humeral complex partially dark brown partially yellow; laterotergites, sternites and hypopygium brown to dark brown [Costa Rica. BIN BOLD:AAL2287]	D. rexhamiltoni Fernandez-Triana & Boudreault, sp. nov.
78(66)	T1 comparatively broader, covering most of dorsal surface of tergum, T1 median length < 1.2× (rarely 1.3×) its width at posterior margin	79
–	T1 comparatively narrower, almost always covering only part of dorsal surface of tergum, T1 median length > 1.5× its width at posterior margin (usually much more)	88
79(78)	Face rostriformis (malar space longer than mandible width) and with slightly elongate mouth parts; T1 and T2 entirely smooth and shiny; T1 very broad at posterior margin, almost quadrate, its length 1.05× its width at posterior margin; ovipositor sheath 1.5× metatibia length; fore wing veins mostly transparent; and pterostigma mostly yellow but with brown margins; and palpi, tegula and humeral complex dark brown [Chile]	D. aceituno Fernandez-Triana & Boudreault, sp. nov.
–	Face almost always not rostriformis (malar space approx. same length than mandible width) and almost always without elongate mouth parts; T1 and T2 from entire to partially sculptured; T1 width variable but usually its length > 1.1× its width at posterior margin; ovipositor sheath length variable but usually < 1.5× metatibia length; fore wing veins color variable; pterostigma usually brown or mostly brown with pale spot at base (rarely strongly yellow-white with very thin brown margins); palpi, tegula and humeral complex variously colored but never the three dark brown (if very rarely pterostigma mostly yellow but with brown margins and palpi, tegula and humeral complex dark brown then T1 and T2 mostly sculptured)	80
80(79)	Comparatively darker colored species, with all legs entirely dark brown to black (except for yellow-brown on anterior 0.1 and/or posterior 0.1 of femur, anterior 0.1–0.3 of metatibia, and sometimes part of trochantellus); pterostigma mostly yellow or strong yellow-white, with brown margins	81
–	Paler colored species, legs always with some segments pale (yellow); pterostigma entirely brown or mostly brown with pale spot at base	82
81(80)	Palpi, tegula and humeral complex dark brown; most veins in fore wing brown; pterostigma mostly yellow with brown margins; comparatively larger species, body length 3.20–3.50 mm, fore wing length 3.50–3.80 mm [Ecuador, 4,000–4,100 m]	D. virgendelparamo Fernandez-Triana & Boudreault, sp. nov.
–	Palpi yellow, tegula pale yellow-brown, humeral complex half yellow and half brown; most veins in fore wing yellow-white; pterostigma strong yellow-white with very thin brown margins; body length 2.80 mm, fore wing length 3.10 mm [Colombia, 2,900 m]	D. lacochaparamo Fernandez-Triana & Boudreault, sp. nov.
82(80)	Ovipositor sheath slightly shorter than metatibia length	83
–	Ovipositor sheath clearly longer (1.15–1.25×) than metatibia length	84
83(82)	F15 cubic, its length 1.1× its width; T2 mostly sculptured, except for central area; tegula yellow; mesosternum entire black [Costa Rica. Host: Telphusa BioLep476 (Gelechiidae)]	D. weaversway Fernandez-Triana & Boudreault, sp. nov.
–	F15 rectangular, its length 1.6× its width; T2 mostly smooth, except along posterior margin; tegula dark brown; mesosternum black but with contrasting pale brown stripe [Costa Rica. Gregarious, host: Microsca hedialis (Thyrididae). BIN BOLD:AAM1098]	D. beryllacosteae Fernandez-Triana & Boudreault, sp. nov.
84(82)	T1 mostly smooth and shiny (sculpture limited along lateral margins on posterior 0.5); overall body comparatively mostly smooth and shiny; body color comparatively paler colored, mostly yellow-brown or reddish brown, including pale brown antenna; trochantelli and tegula yellow, humeral complex partially yellow and partially brown [Costa Rica. Solitary, host: Banisia myrsusalisDHJ01 (Thyrididae). BIN BOLD:AAL2285]	D. alixhamiltonae Fernandez-Triana & Boudreault, sp. nov.
–	T1 mostly sculptured on posterior 0.5 or more; overall body comparatively less smooth and shiny; body color comparatively darker colored, mostly dark brown to black, including dark brown to black antenna; trochantelli, tegula and humeral complex variable but not as above	85
85(84)	Tegula and humeral complex yellow; meso- and metatrochantelli dark brown to black	86
–	Tegula and humeral complex dark brown; trochantelli mostly yellow to yellow-brown	87
86(85)	Body entirely black; T1 slightly broadening towards posterior margin, 1.1–1.2× as long as its width at posterior margin; comparatively larger species, body length 2.80–2.90 mm, fore wing length 3.10–3.30 mm [Costa Rica. Solitary, host: Gelechiidae. BIN BOLD:AAT8840]	D. luishamiltoni Fernandez-Triana & Boudreault, sp. nov.
–	Body mostly dark brown to pale brown; T1 parallel-sided, 1.3× as long as its width at posterior margin; comparatively smaller species, body length 2.30 mm, fore wing length 2.60 mm [Brazil]	D. encruzilhada Fernandez-Triana & Boudreault, sp. nov.
87(85)	Fore wing venation mostly dark brown; pterostigma with small pale yellow-brown spot on anterior 0.1 which is poorly defined; posterior 0.5 of propodeum (beyond transverse carinae of areola) mostly smooth; T2 mostly smooth [Costa Rica. BIN BOLD:AAM5849]	D. antjevirkusae Fernandez-Triana & Boudreault, sp. nov.
–	Fore wing venation mostly pale brown to yellow-brown; pterostigma with comparatively large bright yellow-white spot on anterior 0.2 which is clearly defined; posterior 0.5 of propodeum (beyond transverse carinae of areola) mostly striated; T2 mostly sculptured [Costa Rica. Gregarious, hosts: Anadasmus Janzen25, Stenoma Janzen27 (Depressariidae). BIN BOLD:AAK2061]	D. fredhicksi Fernandez-Triana & Boudreault, sp. nov.
88(78)	Comparatively darker colored species, with all legs entirely dark brown to black (except for yellow-brown on posterior 0.1 of pro- and mesofemora and anterior 0.1–0.2 of tibiae); palpi, tegula and humeral complex dark brown	89
–	Paler colored species, legs always with some segments pale (yellow); coloration of palpi, tegula and humeral complex variable but never the three dark brown	90
89(88)	T1 very strongly narrowing near posterior margin (its length 4.5× its width at posterior margin, and width at anterior margin 3.0× width at posterior margin); propodeum with almost no traces of areola, with only some small, poorly defined carinae from nucha; vein R1 shorter than pterostigma length and approx. as long as than distance between its end and end of vein 3RSb; pterostigma comparatively narrow (3.0× as long as wide) and often with lower anterior margin angulated so that it looks as having four sides; vein r arising from apical 0.7 of pterostigma [Male specimens with some variation in venation, shape of T1 and shape and sculpture of T2] [Ecuador, Peru]	D. rubymacpearsae Fernandez-Triana & Boudreault, sp. nov.
–	T1 narrowing near posterior margin but not as strongly (its length 3.0× its width at posterior margin, and width at anterior margin 2.0× width at posterior margin); propodeum with areola defined on posterior 0.5; vein R1 longer than pterostigma length and much longer than distance between its end and end of vein 3RSb; pterostigma broader and not angulated at lower anterior margin; vein r usually arising at ~ 0.5 of pterostigma length [Chile. BIN BOLD:AAH1316]	D. alerce Fernandez-Triana & Boudreault, sp. nov.
90(88)	T1 strongly narrowing near posterior margin (T1 length ~ 4.0× its width at posterior margin); *and* following areas entirely to mostly smooth: scutellar disc, most of propodeum, T1 and T2	91
–	T1 not strongly narrowing near posterior margin (usually T1 length < 2.5× its width at posterior margin, often much less), if rarely T1 ~ 3.0× its width at posterior margin then T1 evenly narrowing from anterior to posterior margin or narrowing on posterior 0.3 only; and some or all of the following areas not smooth but variably sculptured: scutellar disc, propodeum, T1 and T2	93
91(90)	Propodeum with carinae clearly defining a more or less complete areola; ovipositor sheath comparatively much shorter, its length 0.5× metatibia length; T1 evenly narrowing towards posterior margin (T1 width at anterior margin 2.2× T1 width at posterior margin); legs mostly yellow or pale brown [Costa Rica]	D. carlosviquezi Fernandez-Triana & Boudreault, sp. nov.
–	Propodeum with short carinae weakly defining an areola only on posterior half or less; ovipositor sheath comparatively much longer, ≥ 0.75× metatibia length; T1 strongly to very strongly narrowing near posterior margin; most legs dark brown to black	92
92(91)	F15 comparatively much longer, length/width 1.65–1.75×; ovipositor sheath comparatively shorter, its length 0.75–0.80× metatibia length; T1 comparatively thinner, its length 3.0× or more its maximum width, and T1 width at anterior margin 3.0 × T1 width at posterior margin [Costa Rica, Saint Vincent, Trinidad & Tobago. Solitary, hosts: Antaeotricha spp. (Depressariidae). BIN BOLD:AAN2497]	D. alanflemingi Fernandez-Triana & Boudreault, sp. nov.
–	F15 comparatively much shorter, sub-cubic, length/width 1.15×; ovipositor sheath comparatively longer, its length 1.15× metatibia length; T1 comparatively thicker, its length 2.0× its maximum width, and T1 width at anterior margin 2.0× T1 width at posterior margin [Costa Rica. BIN BOLD:AAD5258]	D. annychaverae Fernandez-Triana & Boudreault, sp. nov.
93(90)	T1 evenly narrowing from anterior to posterior margin and ~ 3.0× its width at posterior margin; *and* ovipositor sheath very short, < 0.55× metatibia length [Costa Rica. BIN BOLD:AAM5852]	D. mehdirheljari Fernandez-Triana & Boudreault, sp. nov.
–	T1 more or less parallel-sided or only narrowing on posterior 0.3 and 2.0× or less its width at posterior margin and/or ovipositor sheath longer than 0.6× metatibia length	94
94(93)	Comparatively darker colored species, with tegula and humeral complex dark brown, all legs brown to dark brown (except sometimes for yellow protibia), tergites dark brown to black, all sternites and hypopygium dark brown; and T2 mostly covered by weak sculptured; and ovipositor sheath length ~ 1.2× metatibia length; and T1 mostly sculptured and parallel-sided but narrowing on posterior 0.3 [Chile]	D. nothofagus Fernandez-Triana & Boudreault, sp. nov.
–	Comparatively paler colored species, with at least some body areas yellow or yellow-orange; and/or T2 smooth or mostly sculptured with longitudinal striae; and/or ovipositor sheath length different and/or T1 shape and sculpture often different than above	95
95(94)	Ovipositor sheath clearly longer than metatibia length (≥ 1.25×, usually more)	96
–	Ovipositor sheath clearly shorter or approx. same length than metatibia (0.60–1.10×)	106
96(95)	Sculpture of T1 and T2 different, T1 mostly sculptured with longitudinal striae on posterior 0.5–0.6 (but sometimes with central smooth area near posterior margin) and T2 entirely smooth [Brazil]	D. robmacewani Fernandez-Triana & Boudreault, sp. nov.
–	Sculpture of T1 and T2 similar, *either* T1 and T2 mostly to entirely smooth or T1 and T2 mostly sculptured	97
97(96)	T1 mostly smooth, at most with weak sculpture on posterior 0.3; T2 smooth	98
–	T1 mostly sculptured with longitudinal striae; T2 entirely to mostly sculptured (at most with small, smooth area centrally)	101
98(97)	Ovipositor tip apically sinuate; T2 smooth [Costa Rica. BIN BOLD:AAM5750]	D. kasiiya Fernandez-Triana & Boudreault, sp. nov.
–	Ovipositor tip not sinuate, if with tip weakly sinuate, then T2 strongly sculptured	99
99(98)	Propodeum with complete areola; tegula and humeral complex white or yellow; pterostigma much paler colored, either mostly yellow-white with thin brown margins, or very pale brown (almost transparent) with small, paler spot at base	100
–	Propodeum without areola, with some striae on posterior margin near nucha; tegula and humeral complex dark brown; pterostigma mostly dark brown with pale spot at base [Canada, Cuba, United States. Solitary, host: Homoeosoma electellum (Pyralidae). Partial sequences available]	D. homoeosomae (Muesebeck, 1933)
100(99)	Ovipositor sheath shorter than metasoma length and 1.3× as long metatibia length; anteromesoscutum coarsely punctured; T1 with some weak sculpture on posterior 0.5, especially along margins [Costa Rica. Solitary, host: Antaeotricha Janzen221 (Depressariidae). BIN BOLD:AAY4695]	D. kenzabaddouae Fernandez-Triana & Boudreault, sp. nov.
–	Ovipositor sheath longer than metasoma length and 1.6–1.7× as long as metatibia length; anteromesoscutum very finely punctate anteriorly, smooth and shiny posteriorly; T1 smooth [Colombia, Dominican Republic, Puerto Rico, Saint Vincent, Trinidad & Tobago]	D. politiventris Muesebeck, 1958
101(97)	T2 with central smooth area; T2 width at posterior margin < 3.0× its central length	102
–	T2 entirely sculptured; T2 width at posterior margin > 3.5× (usually > 4.0×) its central length	103
102(101)	Tegula black, humeral complex dark brown; profemur and protibia mostly brown to dark brown; all trochantelli dark brown to black; pterostigma entirely brown [Costa Rica. BIN BOLD:AAM5739]	D. dole Fernandez-Triana & Boudreault, sp. nov.
–	Tegula yellow, humeral complex half yellow half brown; profemur and protibia yellow; pro- and mesotrochantelli yellow; pterostigma with pale spot at base [Ecuador]	D. puyo Fernandez-Triana & Boudreault, sp. nov.
103(101)	Ovipositor tip weakly sinuate; pterostigma with pale spot very small, 0.1 pterostigma length [Costa Rica. BIN BOLD:AAX8664]	D. rudyamadori Fernandez-Triana & Boudreault, sp. nov.
–	Ovipositor tip slightly and evenly curving downwards but not sinuate; pterostigma pale spot 0.2–0.3 pterostigma length	104
104(103)	All trochantelli yellow; profemur entirely and mesofemur mostly (except for longitudinal brown bands on margins) yellow [Costa Rica. Solitary, host: Dichomeris designatellaDHJ04 (Gelechiidae). BIN BOLD:AAI6323]	D. susanabramsae Fernandez-Triana & Boudreault, sp. nov.
–	All trochantelli dark brown; anterior 0.3–0.4 of profemur and entire mesofemur brown	105
105(104)	Most veins transparent to yellow white; pterostigma comparatively elongate, its length > 3.0× its maximum height [Costa Rica, Mexico. BIN BOLD:AAX8653]	D. moniquegilbertae Fernandez-Triana & Boudreault, sp. nov.
–	Most veins yellow-brown or brown; pterostigma comparatively thicker, its length ~ 2.0× its maximum width [United States. Solitary, hosts: Dioryctria spp, Rhyacionia spp. (Pyralidae, Tortricidae)]	D. bushnelli (Muesebeck, 1933)
106(95)	Comparatively darker colored species, with tegula and humeral complex dark brown, all legs brown to dark brown (except for yellow protibia), tergites dark brown to black, all sternites and hypopygium dark brown [Peru]	D. machupichu Fernandez-Triana & Boudreault, sp. nov.
–	Comparatively paler colored species, with at least some body areas yellow or yellow-orange	107
107(106)	Scutellar disc mostly with shallow punctures; and T1 relatively thinner, with median length 2.0–2.2× its width at posterior margin; and T1 with central hump; and tegula and humeral complex yellow; and mesofemur, metatibia (except for small pale spot on anterior 0.1) and metatarsus dark brown [Costa Rica. BIN BOLD:AAI9747]	D. jasonkelleyi Fernandez-Triana & Boudreault, sp. nov.
–	Scutellar disc smooth and shiny, without punctures; either with T1 relatively thicker, with median length < 2.0× its width at posterior margin; and/or T1 without central hump; and/or tegula and humeral complex darker; and/or mesofemur, metatibia and metatarsus partially or entirely pale colored	108
108(107)	Pterostigma pale (yellow or white-yellow), at most with thin margins which are slightly darker	109
–	Pterostigma mostly dark (brown), at most with small pale spot on anterior 0.1 or, very rarely, with small spot centrally which is slightly paler than rest of pterostigma	111
109(108)	Tegula and humeral complex yellow; anteromesoscutum with punctures on most of its surface and scutellar disc with some punctures [Costa Rica. BIN BOLD:AAE8602]	D. josephfridmani Fernandez-Triana & Boudreault, sp. nov.
–	Tegula and humeral complex dark brown; anteromesoscutum and scutellar disc mostly smooth and shiny	110
110(109)	Pterostigma bright yellow-white; T2 entirely smooth; T1 mostly smooth, with weak punctures centrally; fore wing vein R1 longer than pterostigma and > 3.5× as long as the space between its end and end of vein 3RSb [Chile]	D. moniqueae Fernandez-Triana & Boudreault, sp. nov.
–	Pterostigma pale yellow-brown but with thin brown margins; T2 mostly sculptured, with small area smooth along posterior margin; T1 rugose on posterior half; fore wing vein R1 around same length (sometimes shorter) than pterostigma and 3.0× or less as long as the space between its end and end of vein 3RSb [Algeria, Chile, Colombia, Peru, Spain, Venezuela. Solitary, hosts: Phthorimaea operculella, Keiferia lycopersicella, Tuta absoluta (Gelechiidae)]	D. gelechiidivoris (Marsh, 1975)
111(108)	Metatibia mostly yellow (except for dark brown spot on posterior 0.1–0.2); part of metafemur (anterior 0.1–0.2 and posterior 0.1) yellow	112
–	Metatibia ≥ 0.5–0.8 dark brown; metafemur usually almost entirely dark brown to black	113
112(111)	T1 comparatively wider, its median length 1.25× its width at posterior margin; humeral complex yellow, same color than tegula; fore wing vein 1CU medially raised or arched in a sharp angle [Costa Rica. Solitary, hosts: Gonionota Janzen22 (Depressariidae), unidentified Gelechiidae with interim name ‘gelJanzen01 Janzen23’. BIN BOLD:AAI9755]	D. christinaagapakisae Fernandez-Triana & Boudreault, sp. nov.
–	T1 comparatively thinner, its median length 1.65× its width at posterior margin; humeral complex almost entirely brown, clearly darker than yellow tegula; fore wing vein 1CU straight [Costa Rica, French Guiana. Solitary, host: unidentified Depressariidae. BIN BOLD:ACI3397]	D. sigifredomarini Fernandez-Triana & Boudreault, sp. nov.
113(111)	Tegula yellow	114
–	Tegula dark brown to black	119
114(113)	Humeral complex yellow; T2 mostly to entirely smooth and comparatively very transverse, its width at posterior margin 4.0× its central length; T1 mostly smooth (only with fine sculpture laterally near posterior margin) [Caribbean species]	115
–	Humeral complex mostly to entirely brown; T2 mostly sculptured (at least around margins) and comparatively less transverse, its width at posterior margin 2.2–3.0× its central length; T1 with some sculpture, sometimes strong on posterior half [Central and South America species]	116
115(114)	Propodeum mostly smooth and without defined areola; pterostigma entirely brown; mesoscutum shiny and with extremely shallow and separate punctures; lunules semicircular and comparatively short; fore wing with vein r arising well beyond middle of pterostigma; metafemur mostly yellow-brown (sometimes darker on upper and lower margins), metatibia darker on posterior 0.2 or less; body color, including coxae, mostly black [Puerto Rico. Probably gregarious, hosts: Maruca vitrata, Omiodes indicata (Pyralidae)]	D. hedyleptae (Muesebeck, 1958)
–	Propodeum mostly with punctures and rugulosities and with complete and strongly defined areola; pterostigma with pale spot on anterior 0.3; lunules triangular and comparatively very high; fore wing with vein r arising around middle of pterostigma; metafemur and posterior 0.5 of metatibia dark reddish brown; body color, including coxae, mostly dark reddish brown [Trinidad & Tobago. Solitary, host: Oiketicus kirbyi (Psychidae)]	D. oiketicus Fernandez-Triana & Boudreault, sp. nov.
116(114)	T1 2.2× as long as wide at posterior margin [Costa Rica. BIN BOLD:AAI9746]	D. alejandromarini Fernandez-Triana & Boudreault, sp. nov.
–	T1 1.3–1.7× as long as wide at posterior margin	117
117(116)	Ovipositor sheath clearly shorter than metatibia length (0.7×) [Costa Rica. Solitary, host: unidentified Pyralidae. BIN BOLD:ABX5620]	D. luzmariaromeroae Fernandez-Triana & Boudreault, sp. nov.
–	Ovipositor sheath as long as metatibia or longer	118
118(117)	Anteromesoscutum mostly with rather coarse punctures; propodeum with almost complete areola (open anteriorly); ocelli comparatively smaller, ocular ocellar line > 3.0× diameter of posterior ocellus; pterostigma mostly dark brown (with pale spot on anterior 0.1 or less) [Costa Rica. Solitary, hosts: Olethreutes spp. (Tortricidae). BIN BOLD:AAE8612]	D. anacamposae Fernandez-Triana & Boudreault, sp. nov.
–	Anteromesoscutum mostly shiny, punctures sparse, superficial and fading towards posterior margin; propodeum without defined areola but with faint U-shaped depression medially; ocelli comparatively larger, ocular ocellar line ~ 2.0× diameter of posterior ocellus; pterostigma mostly pale brown (with pale spot anteriorly) [Chile, Juan Fernández islands]	D. evadne (Nixon, 1955)
119(113)	T2 entirely to mostly smooth (if there is some sculpture, it is limited to margins of tergite)	120
–	T2 entirely to mostly sculptured	122
120(119)	Gena with white spot; smaller size, body length and fore wing length 1.80 mm; propodeum areola open anteriorly, propodeum generally with relatively few sculptures or carinae [Costa Rica. BIN BOLD:AAM5853]	D. puschendorfi Fernandez-Triana & Boudreault, sp. nov.
–	Gena without white spot; larger size, body length and fore wing length 2.40–2.60 mm; propodeum areola complete, propodeum generally sculptured and with well-defined carinae	121
121(120)	Comparatively darker colored specimen, with antenna pedicel, most of mesofemur and entire mesosternum dark brown to black; propodeum mostly sculptured, with lateral striation on most areas between carinae; anteromesoscutum mostly covered with relatively coarse punctures; T1 slightly narrowing near posterior margin; T2 smooth [Costa Rica. BIN BOLD:AAJ1390]	D. timrichi Fernandez-Triana & Boudreault, sp. nov.
–	Comparatively paler colored specimen, with pedicel, most of mesofemur and longitudinal band on mesosternum yellow to yellow-brown; propodeum mostly smooth, with few areas between carinae with some striation; anteromesoscutum punctures less coarse on posterior 0.3–0.4; T1 slightly widening near posterior margin; T2 usually with some sculpture [Costa Rica. Gregarious, hosts: Microsca spp. (Thyrididae). BIN BOLD:ABX6008]	D. sarahoconnorae Fernandez-Triana & Boudreault, sp. nov.
122(119)	Ovipositor sheath distinctly shorter than metatibia length (< 0.7×)	123
–	Ovipositor sheath slightly longer (1.1–1.2×) than metatibia length	124
123(122)	T1 clearly narrowing towards posterior margin (T1 width at anterior margin 1.2 × T1 width at posterior margin) and comparatively less broad (T1 length 2.2 × T1 width at posterior margin); T2 trapezoidal, its width at posterior margin ~ 2.0× its central length; propodeum areola less defined anteriorly; profemur brown on anterior ~ 0.5; metatibia with anterior 0.2 yellow-white, posterior 0.8 dark brown [Costa Rica. BIN BOLD:AAM5851]	D. jessiehillae Fernandez-Triana & Boudreault, sp. nov.
–	T1 parallel-sided (same T1 width at anterior and posterior margins) and comparatively much broader (T1 length 1.2 × T1 width at posterior margin); T2 transverse, its width at posterior margin > 3.0× its central length; propodeum areola well defined and bounded by carinae; profemur entirely yellow; metatibia with anterior 0.4 yellow, posterior 0.6 brown [Costa Rica. Solitary, host: unidentified Gelechiidae. BIN BOLD:ACB1629]	D. stevestroudi Fernandez-Triana & Boudreault, sp. nov.
124(122)	Anterior 0.5 of profemur and entire mesofemur brown; and antenna shorter than body length; and comparatively smaller size, body length 2.10 mm, fore wing length 2.30 mm [Brazil]	D. stephmae Fernandez-Triana & Boudreault, sp. nov.
–	Pro- and mesofemora usually entirely yellow (if rarely femora darker, then antenna much longer than body and comparatively larger size, body length 2.60 mm, fore wing length 2.75 mm); antenna usually as long as body length or longer; usually species larger than 2.00 mm	125
125(124)	Anterior 0.5 of profemur and entire mesofemur brown; ovipositor sheath distinctly (> 1.2×) longer than metatibia length [Costa Rica. Solitary, hosts: Chlamydastis vividella, Stenoma Janzen199 (Depressariidae). BIN BOLD:ACC1295]	D. bradzlotnicki Fernandez-Triana & Boudreault, sp. nov.
–	Pro- and mesofemora entirely yellow; ovipositor sheath from slightly longer (1.1× or less) to distinctly shorter (< 0.8×) than metatibia length	126
126(125)	Comparatively longer F15, its length > 1.5× its width; usually body comparatively less shiny and less smooth; sculpture on anteromesoscutum and propodeum coarser and more deeply indicated; comparatively darker colored, with paler areas (such as longitudinal strip on metasternum and most of first pair of legs) yellow to yellow-brown [Costa Rica. Solitary, host: Microsca paullula (Thyrididae). BIN BOLD:AAC2174]	D. scottmilleri Fernandez-Triana & Boudreault, sp. nov.
–	Comparatively shorter, cubic, F15, its length ≤ 1.1× its width; body comparatively shinier and smoother; sculpture on anteromesoscutum and propodeum shallower; comparatively paler colored with some areas (such as longitudinal strip on metasternum and most of first pair of legs) bright yellow to white-yellow [Costa Rica. Gregarious, host: Collinsa ferreiceps (Thyrididae)]	D. robpringlei Fernandez-Triana & Boudreault, sp. nov.

Taxonomic treatment of species, in alphabetical order

Dolichogenidea aceituno Fernandez-Triana & Boudreault, sp. nov.

https://zoobank.org/1930F0BD-E5B4-4F6E-823C-52166B8B060D

Fig. 6A–H

Type material.

Holotype. Chile • Female, CNC; Atacama, Chanaral de Aceituno; 12.x.1958; L. E. Pena leg.; Voucher code: CNC1196525. Paratypes. Chile • 3 Males, CNC; CNC1196544, CNC1196545, CNC1196567.

Figure 6.

Dolichogenidea aceituno Fernandez-Triana & Boudreault holotype female CNC1196525 A head, fronto-lateral B habitus, lateral C head, frontal D head, dorsal E wings F propodeum & T1–T3, dorsal G metasoma, dorsal H mesosoma, dorsal.

Diagnostic description.

Face rostriformis (malar space longer than mandible width) and with slightly elongate mouth parts; T1 and T2 entirely smooth and shiny; T1 very broad at posterior margin, almost quadrate, its length 1.05× its width at posterior margin; ovipositor sheath 1.5× metatibia length; fore wing veins mostly transparent; pterostigma mostly yellow but with brown margins; palpi, tegula and humeral complex dark brown; all coxae black to dark brown; profemur dark brown on posterior half, meso- and metafemora entirely dark brown; body length: 3.70 mm; fore wing length: 3.90 mm. Among all species with smooth T1 and T2, D. aceituno is very distinctive because of its elongate face and mouth parts, dark coloration of coxae and femora and broad T1.

Distribution.

Chile.

Biology.

No host data available.

DNA barcoding data.

No data.

Etymology.

Named after the type locality, the Chañaral de Aceituno, in the Chanaral Island (Isla Chanaral) where the Pingüino de Humboldt National Reserve, an important nature reserve for marine wildlife, is located.

Dolichogenidea acrobasidis (Muesebeck), 1921

Figs 7A–H, 8A, B

Notes.

This species has not been found in the Neotropical region, but it is included in the key above because of its occurrence in southern USA states (Florida and Mississippi).

Figure 7.

Dolichogenidea acrobasidis (Muesebeck) female CNCHYM00971 A habitus, lateral B head, frontal C wings D head & mesoscutum, dorsal E ovipositor F propodeum & metasoma, dorsal G metasoma, dorsal H propodeum & T1–T3, dorsal.

Figure 8.

Dolichogenidea acrobasidis (Muesebeck) holotype female A habitus, dorsal B habitus, lateral.