Research Article |
Corresponding author: Sarah Meierotto ( s.meierotto@uky.edu ) Corresponding author: Michael J. Sharkey ( msharkey@uky.edu ) Academic editor: Dominique Zimmermann
© 2019 Sarah Meierotto, Michael J. Sharkey, Daniel H. Janzen, Winnie Hallwachs, Paul D. N. Hebert, Eric G. Chapman, M. Alex Smith.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Meierotto S, Sharkey MJ, Janzen DH, Hallwachs W, Hebert PDN, Chapman EG, Smith MA (2019) A revolutionary protocol to describe understudied hyperdiverse taxa and overcome the taxonomic impediment. Deutsche Entomologische Zeitschrift 66(2): 119-145. https://doi.org/10.3897/dez.66.34683
|
Here we elucidate and justify a DNA barcode approach to insect species description that can be applied to name tens of thousands of species of Ichneumonoidea and many other species-rich taxa. Each description consists of a lateral habitus image of the specimen, a COI barcode diagnosis, and the holotype specimen information required by the International Code of Zoological Nomenclature. We believe this approach, or a slight modification of it, will be useful for many other underdescribed hyperdiverse taxa, especially in the tropics. Due to the extreme species-richness of the Ichneumonoidea, the very low percentage of described species, and the lack of detailed biological information for most described species, the standard taxonomic approach is inefficient and overwhelmingly time consuming. A DNA barcode-based approach to initial description will provide a solid foundation of species hypotheses from which more comprehensive descriptions can be developed as other data, time, and budgets permit. Here we elucidate this view and detailed methodology that can generally be applied to species-rich underdescribed taxa. A real example is given by describing species in two genera, Hemichoma and Zelomorpha, reared from the Área de Conservación Guanacaste in northwestern Costa Rica. The generic type species Zelomorpha arizonensis is given a DNA barcode diagnosis and the following new species are described: Zelomorpha angelsolisi, Zelomorpha bobandersoni, Zelomorpha danjohnsoni, Zelomorpha donwindsori, Zelomorpha effugia, Zelomorpha johnchemsaki, Zelomorpha kellyanneae, Zelomorpha larrykirkendalli, Zelomorpha mariyavladmirovnae, Zelomorpha mikeiviei, Zelomorpha myricagaleae, Zelomorpha noahjaneae, Zelomorpha paulgoldsteini, Zelomorpha terryerwini, Zelomorpha willsflowersi, Hemichoma donwhiteheadi, Hemichoma frankhovorei, and Hemichoma johnkingsolveri.
Agathidinae, barcode, Braconidae, COI, Hemichoma, Hymenoptera, Ichneumonoidea, Zelomorpha
Systematists have many powerful tools at their disposal for discovering, delimiting and describing new species, and an integrated taxonomic approach, combining morphological characters, identification keys, phylogenetic analyses with multiple molecular markers, and ecological data, is currently the gold standard for quality descriptions of new species (
We propose the description of new species based primarily on the DNA barcode molecular marker as a first step in the systematic study of terminal taxa in the highly diverse superfamily Ichneumonoidea. These descriptions are meant to encourage and accelerate 1) the accumulation of additional information on the described species, 2) scientific discussion of the groups treated, 3) opportunities for the refinement of presented species hypotheses as well as those long believed to be established (e.g.,
The superfamily Ichneumonoidea contains the two most species-rich families of Hymenoptera, i.e., Braconidae and Ichneumonidae. As parasitoids, ichneumonoids exert strong top-down control on their hosts, contributing to ecosystem stability and diversity (
The Ichneumonoidea contains over 44,000 valid species as of 2016 (
Recent revisions of ichneumonoids in the subfamilies Agathidinae and Microgastrinae have investigated the utility of the DNA barcoding region of the gene cytochrome c oxidase subunit I (COI) for species delimitation, paired with morphological and ecological host-use characters (e.g.,
While there have been some calls to use molecular species descriptions (
Most specimens were collected by rearing host caterpillars in Área de Conservación Guanacaste (ACG) in northwestern Costa Rica (
Molecular work was carried out at the Centre for Biodiversity Genomics (CBG) using their standard protocols. A leg of each specimen was destructively sampled for DNA extraction using a glass fiber protocol (
Sequences (barcodes) were assigned to operational taxonomic units called barcode index numbers (BINs) that were generated in tandem with the neighbor-joining tree of the barcodes themselves (Appendix
Morphology and host information were compared to BIN assignments that package the NJ tree. Specimen groupings, a BIN, suggested by all data sources were considered to be species. However, for other higher taxa there are cases where several obvious species are packaged into one BIN (e.g.,
Consensus barcodes were created for each species using BioEdit (
For Zelomorpha arizonensis, the only previously described species that we included in the dataset, a number of specimens collected in the southwestern USA and Mexico were included in the NJ tree in addition to the sequences of specimens from ACG.
Holotypes are deposited in the insect collection in the Biology Department of Utah State University in Logan, Utah (
Some host species are still awaiting full identification and are given interim names, generally based on their barcodes, just as is the case with the wasps described here. For example, Hemiceras plusiataDHJ01 is identified to the genus Hemiceras by classical morphology-based criteria. However, it is one of the two reared in a species complex that used to be known as H. plusiata. One of the two is the actual H. plusiata, and the other is new, but which is which cannot be determined until the holotype is barcoded, or more closely examined morphologically. This was the case with Udranomia kikkawaiDHJ01, H. kikkawaiDHJ02 and H. kikkawaiDHJ03, recently rendered into three species (
Three hundred thirty-six specimens of Zelomorpha and Hemichoma with COI barcodes were determined to represent 20 species in two genera by their barcodes, their BINs, and by concomitant morphological inspection. BIN assignments were the same as final species hypotheses (Appendix
Zelomorpha arizonensis (by monotypy) (
Zelomorpha can be distinguished from all other Agathidinae genera by the following combination of morphological characters: fore tarsal claws cleft and not pectinate; foretibial spur shorter than first tarsomere; ovipositor shorter than half the length of the metasoma; frons bordered by carinae; hind trochantellus with one or two longitudinal ridges; notauli variable, usually distinct; gena not elongate.
The species of Zelomorpha are koinobiont solitary endoparasitoids of free-living, late instar medium-small Lepidoptera larvae (
Zelomorpha occur only in the New World, from the southern USA to Argentina and are primarily Neotropical (
Including the fifteen new species described here, there are 67 described species of Zelomorpha (
Nucleotides 43–45 TTA, 54–57 CTTT, 75 G, 136–138 GTG, 165 T, 321 G, 417 G, 462 G, 477 C, 561 G, 684 G.
This species has characteristics associated with nocturnal habits: pale coloration, large compound eyes and ocelli. Specimens were reared from caterpillars in the family Erebidae feeding on Fabaceae: Azeta ceramina on Acosmium panamense, Chabora repugnalisDHJ01 on Indigofera costaricensis, and Coenipeta bibitrix on Enterolobium cyclocarpum. Host caterpillars were collected in April, May, and November.
Holotype ♀: DHJPAR0009310 (ASBR577-06), Costa Rica, Área de Conservación Guanacaste, Sector Mundo Nuevo, Punta Plancha, GPS: 10.7416, -85.42734, 420 m elevation, Mariano Pereira coll., reared from Azeta ceramina 05-SRNP-56517, host collected 30 May 2005, wasp eclosed 17 June 2005, (
Zelomorpha angelsolisi is named in honor of Angel Solis of INBio and the Museo Nacional de Costa Rica, a master taxonomist of Coleoptera and curator who has massively contributed to the inventory of Costa Rican Coleoptera.
Nucleotides 515 C, 648 T
Adults of this species have characteristics associated with nocturnal habits: pale coloration, large compound eyes and large ocelli. All individuals were reared from Bulia mexicana (Erebidae) caterpillars feeding on mature leaves of Prosopis juliflora (Fabaceae) at the edge of ACG mangrove swamps in the month of July.
The host of Z. arizonensis from the type locality in the southwestern United States is unknown. However, the range of Prosopis juliflora extends northwards through Mexico and into the United States, where it is fed upon by several species of Bulia. P-distances between specimens from Costa Rica and the US are close to 1.5% (Fig.
Figured specimen ♀: DHJPAR0052709 (ASHYM2063-13), Costa Rica, Área de Conservación Guanacaste, Sector Santa Rosa, Argelia, GPDS: 10.78004, -85.66405, 5 m elevation, Guillermo Pereira coll., reared from Bulia mexicana 13-SRNP-17758, host collected 13 July 2013, wasp eclosed 29 July 2013, (
Zelomorpha arizonensis was named for its holotype locality.
Nucleotides: 72–75 GGGT, 163 G, 222–225 GGGG, 264 G
All known individuals were reared from Hemiceras plusiataDHJ01 or Hemiceras plusiataDHJ02 (Notodontidae) feeding on Tachigali costaricense (Fabaceae); the caterpillars are indistinguishable without barcoding them. Host caterpillars were collected in January, February, April, and June through October.
Holotype ♀: DHJPAR0028037 (ASHYE274-08), Costa Rica, Área de Conservación Guanacaste, Sector Pitilla, Estacion Quica, GPS: 10.99697, -85.39666, 470 m elevation, Mauricio Siezar coll., reared from Hemiceras plusiataDHJ01 08-SRNP-71265, host collected 10 July 2008, wasp eclosed 11 August 2008, (
Zelomorpha bobandersoni is named in honor of Bob Anderson of the Canadian Museum of Nature, Ottawa, in recognition of his taxonomic and curatorial support for understanding the Curculionidae of Costa Rica.
Nucleotides: 98 G, 111 G, 264 C, 310 G, 375 A, 452 T, 495 A, 507 G, 513 G, 648 G
Holotype ♀: DHJPAR0009409 (ASBR676-06), Costa Rica, Área de Conservación Guanacaste, Sector Cacao, Quebrada Otilio, GPS: 10.88996, -85.47966, 550m elevation, Dunia Garcia coll., reared from Diastema morata 05-SRNP-45510, host collected 7 June 2005, wasp eclosed 14 July 2005, (
Zelomorpha danjohnsoni is named in honor of C. Dan Johnson (RIP) of Arizona State University, in recognition of his taxonomic support for understanding the Bruchinae (Chrysomelidae) of Costa Rica.
Nucleotides: 78 A, 213 C, 243 A, 390 G, 429 G, 456 G, 506–507 CT, 513 T, 585 G, 588 G, 603 C, 636 C, 660 G, 678–679 TG
The two specimens of this species were reared from caterpillars in the Euteliidae, Paectes lunodes feeding on very young leaves of Ocotea veraguensis (Lauraceae) and Paectes fuscescens feeding on young leaves of the introduced species Anacardium occidentale (Anacardiaceae). Host caterpillars were collected in November and July.
Holotype ♀: DHJPAR0048721 (ACGBA2263-12), Costa Rica, Área de Conservación Guanacaste, Sector El Hacha, Los Almendros, GPS: 11.03226, -85.52776, 290 m elevation, Elieth Cantillano coll., reared from Paectes fuscescens 11-SRNP-23258, host collected 15 November 2011, wasp eclosed 9 January 2012, (
Zelomorpha donwindsori is named in honor of Don Windsor of the Smithsonian Tropical Research Institute in Panama, a master taxonomist in Chrysomelidae who also contributed to the early development of ACG. The timbers from his original house are part of an ACG caterpillar rearing barn.
Nucleotides: 46 A, 96–97 TG, 102 T, 124–127 TTAA, 130 G, 285 G, 352–353 TC
This species has been reared only from Cosmosoma hercyna (Erebidae) caterpillars feeding on mature leaves of Lacistema aggregatum (Lacistemataceae), and Lozania pittieri (Lacistemataceae). Hosts were collected in September, November, January, and February.
Holotype ♀: DHJPAR0015541 (ASAG227-07), Costa Rica, Área de Conservación Guanacaste, Sector Rincon Rain Forest, Vochysia, GPS: 10.86666, -85.24528, 320 m elevation, Minor Carmona coll., reared from Cosmosoma hercyna 05-SRNP-43568, host collected 30 November 2005, wasp eclosed 27 December 2005, (
Zelomorpha effugia is named in honor of the podcast Escape Pod, whose short science fiction stories provided the first author with inspiration and motivation during the production of this manuscript.
Nucleotides: 261 G, 279 C, 537–538 GC, 571 G
All 24 rearing records for this species are from Hemiceras pallidula (Notodontidae) feeding on mature leaves of Inga vera and Inga oerstediana (Fabaceae). Two of the hosts were collected in October and the others in July.
Members of Z. johnchemsaki are similar to Z. bobandersoni in COI sequence and morphology, but the large samples of the two species show consistent differences in color pattern and host preference, and fall in different BINs. They are an excellent example of a shallow split in an NJ tree that definitely represents two species.
Holotype ♀: DHJPAR0040547 (ASHYE2683-11), Costa Rica, Área de Conservación Guanacaste, Sector Pitilla, Bullas, GPS: 10.9867, -85.38503, 440 m elevation, Ricardo Calero coll., reared from Hemiceras pallidula 09-SRNP-71580, host collected 14 July 2009, wasp eclosed 10 August 2009, (
Zelomorpha johnchemsaki is named in honor of John Chemsak (RIP) of the University of California, Berkeley, in recognition of his taxonomic support for understanding the ACG Cerambycidae and teaching DHJ about them in the 1960’s.
Nucleotides: 348 C, 421 A
Holotype ♀: DHJPAR0015536 (ASAG222-07), Costa Rica, Área de Conservación Guanacaste, Sector Del Oro, Quebrada Raiz, GPS: 11.02865, -85.48669, 280 m elevation, Lucia Ríos coll., reared from Nephodia Janzen18, 05-SRNP-25234, host collected 21 November 2005, wasp eclosed 10 December 2005, (
Zelomorpha kellyanneae is named in honor of Kelly Meierotto, sister of SM and up and coming archaeologist.
Nucleotides: 81 G, 273 G, 324 T, 369 A, 432 G, 522 A, 662 G
This species has been reared from three species of Opisthoxia (Geometridae) feeding on very young leaves of three species of Primulaceae: O. molpadia on Parathesis glabra, O. bella on Ardisia compressa, and O. uncinata on Ardisia auriculata. Caterpillars were collected in February, March, June, July, and September.
Holotype ♀: DHJPAR0015540 (ASAG226-07), Costa Rica, Área de Conservación Guanacaste, Sector San Cristobal, Rio Blanco Abajo, GPS: 10.90037, -85.37254, 500 m elevation, Yessenia Mendoza coll., reared from Opisthoxia bella 04-SRNP-4505, host collected 6 September 2004, wasp eclosed 26 September 2004, (
Zelomorpha larrykirkendalli is named in honor of Larry Kirkendall of the University of Bergen, Norway, in recognition of his intense taxonomic interest in Neotropical Scolytidae and Platypodidae, and now, those of ACG.
Nucleotides: 250 A, 354 G, 462 C, 543 G
The single specimen of this species was reared from Ormetica sicilia (Erebidae) feeding on mature leaves of Inga vera (Fabaceae). Unexpectedly, it appears that this wasp eclosed from the moth pupa rather than from a wasp cocoon spun inside the moth cocoon.
Holotype ♀: DHJPAR0023528 (ASHYM280-08), Costa Rica, Área de Conservación Guanacaste, Sector Mundo Nuevo, GPS: 10.77175, -85.434, 305 m elevation, Jose Cortez coll., reared from Ormetica sicilia 07-SRNP-61364, host collected 28 December 2007, wasp eclosed 14 January 2008, (
Zelomorpha mariyavladmirovnae is named in honor of Mariya Frahm, for her guidance and support to SM.
Nucleotides: 111 C, 411 G, 549 G, 567 G, 661 T
This species has been reared from three unidentified, different species of host feeding on two different host plants: a species of Geometridae feeding on Ruellia inundata (Acanthaceae) and another on Solanum hayesii (Solanaceae), and a species of Erebidae feeding on Colubrina spinosa (Rhamnaceae). Host caterpillars were collected in January and June.
Holotype ♀: DHJPAR0029297 (ASHYE704-09), Costa Rica, Área de Conservación Guanacaste, Sector Pitilla, Pasmompa, GPS: 11.01926, -85.40997, 440 m elevation, Calixto Moraga coll., reared from Erebidae 04-SRNP-30170, host collected 12 January 2004, wasp eclosed 6 February 2004, (
Zelomorpha mikeiviei is named in honor of Mike Ivie of Montana State University, a master taxonomist in Coleoptera of who has massively contributed to the knowledge base of the inventory of Caribbean Coleoptera and ACG biodiversity inventory.
Nucleotides: 44 C, 55 A, 64 G, 98 C, 126 C, 135 G, 163 T, 168 G, 183–186 GGTA, 246 C, 258 G, 357–358 GG, 369 G, 381 C, 400–401 AA, 505 T, 519–520 CG, 525 G, 570 A, 603 G, 606 G
The single specimen of this species was reared from Speocropia Poole01 Noctuidae feeding on mature leaves of Smilax spinosa (Smilacaceae).
Holotype ♀: DHJPAR0028033 (ASHYE270-08), Costa Rica, Área de Conservación Guanacaste, Sector Del Oro, Quebrada Trigal, GPS: 11.02681, -85.49547, 290 m elevation, Lucia Ríos coll., reared from Speocropia Poole01 08-SRNP-21458, host collected 11 June 2008, wasp eclosed 8 July 2008, (
Zelomorpha myricagaleae is named in honor of Myrica Gale Meierotto, cousin of SM and fierce competitor.
Nucleotides: 108 G, 123 G, 333 G, 519 A, 693 CG
Specimens of this species were reared from three species of Euteliidae feeding on young leaves of Anacardiaceae: Paectes fuscescens on introduced Anacardium occidentale, Eutelia chrysotermina on Anacardium excelsum, and Paectes Poole10 on Mosquitoxylum jamaicense. Caterpillars were collected in July and November.
Holotype ♀: DHJPAR0048720 (ACGBA2262-12), Costa Rica, Área de Conservación Guanacaste, Sector El Hacha, Estacion los Almendros, GPS: 11.03226, -85.52776, 290 m elevation, Elieth Cantillano coll., reared from Paectes fuscescens 11-SRNP-23262, host collected 15 November 2011, wasp eclosed 30 December 2011, (
Zelomorpha noahjaneae is named in honor of Noah Jane Meierotto, cousin of SM and an aspiring scientist and possible future entomologist.
Nucleotides: 216 G, 327 G, 345–346 AA, 352–354 ACA, 517 C
This species has been reared from a relatively wide range of hosts in the families Erebidae and Noctuidae, but all hosts feed on leaves of ferns. Caterpillars of paratype specimens were collected in every month except March and April.
Holotype ♀: DHJPAR0040222 (ASHYE2389-11), Costa Rica, Área de Conservación Guanacaste, Sector Del Oro, Quebrada Serrano, GPS: 11.00025, -85.45614, 585 m elevation, Roster Moraga coll., reared from Callopistria mexicana 10-SRNP-21839, host collected 5 August 2010, wasp eclosed 29 August 2010, (
Zelomorpha paulgoldsteini is named in honor of Paul Goldstein of the USDA Systematic Entomology Laboratory at the Smithsonian Institution, in honor of his inordinate fondness for the fern-eating caterpillars parasitized by this wasp.
Nucleotides: 66 G, 359 G, 492 C, 621 G
Holotype ♀: DHJPAR0054486 (ASHYD3651-14), Costa Rica, Área de Conservación Guanacaste, Sector Rincon Rain Forest, Jacobo, GPS: 10.94076, -85.3177, 461 m elevation, Edwin Apu coll., reared from Iscadia Poole02DHJ03 13-SRNP-80618, host collected 13 November 2013, (
Zelomorpha terryerwini is named in honor of Terry Erwin of the Smithsonian Institution, a master taxonomist of Coleoptera who has massively contributed to the inventory of Latin American Coleoptera and pesticide-fogged more trees than any other entomologist.
Hemichoma fenestratum Enderlein, 1920.
Hemichoma shares diagnostic morphological characters with Zelomorpha except: notauli absent, mesoscutum lacking distinct lobes; gena greatly produced posteroventrally.
Members of Hemichoma are, like Zelomorpha, koinobiont endoparasitoids of late instar lepidopteran larvae. The solitary wasp larva emerges from the prepupal larva after it has spun its cocoon, and spins its own cocoon inside the host cocoon next to the cadaver.
Restricted to the New World, known from Mexico to Argentina.
Including the three species described here, there are eight described species of Hemichoma.
Nucleotides: 72 G, 78 G, 90 G, 114 G, 162 T, 168 A, 204 C, 207 G, 216 G, 225 G, 306 G, 318 T, 322 T, 346 G, 357 T, 409–410 GC, 414 G, 492 A, 516 G, 564 A, 585 GC
All specimens of this species were reared from Pelochyta misera (Erebidae: Arctiinae). Food plants include Heliocarpus appendiculatus (Malvaceae), the introduced species Psidium guajava (Myrtaceae), Inga oerstediana, and Erythrina costaricensis (Fabaceae). Host caterpillars were collected in June, August, November, and October.
Holotype ♀: DHJPAR0016918 (ASBR891-07), Costa Rica, Área de Conservación Guanacaste, Sector San Cristobal, Sendero Huerta, GPS: 10.9305, -85.37223, 527 m elevation, Elda Araya coll., reared from Pelochyta misera 06-SRNP-9643, host collected 27 November 2006, (
Hemichoma donwhiteheadi is named in honor of Don Whitehead (RIP) of the Smithsonian Institution, a master weevil taxonomist who helped greatly with the taxonomy of ACG Curculionidae.
Nucleotides: 117 G, 228 C, 243 A, 357 A, 414 A, 477 T, 513 T, 570 A, 615 G, 645 T, 60 A, 663 T
Multiple species of Halysidota (Erebidae; Arctiinae) are the hosts for this wasp: H. orientalis, H. pectenella, H. schausi, and H. underwoodi feeding on mature leaves of Trema micrantha (Cannabaceae), Bernardia nicaraguensis (Euphorbiaceae), and Acalypha macrostachya (Euphorbiaceae). Host caterpillars of type specimens were collected between the months of September and December.
Holotype ♀: DHJPAR0054503 (ASHYD3668-14), Costa Rica, Área de Conservación Guanacaste, Sector Pitilla, Medrano, GPS: 11.01602, -85.38053, 380 m elevation, Ricardo Calero coll., reared from Halysidota schausi 13-SRNP-71924, host collected 2 December 2013, wasp eclosed 12 January 2014, (
Hemichoma frankhovorei is named in honor of Frank Hovore (RIP) of California, a master cerambycid taxonomist who helped greatly with the taxonomic inventory of Costa Rican Cerambycidae.
Nucleotides: 77 C, 84 G, 108 T, 111 A, 122 C, 141 T, 297 T, 327 G, 357 G, 414 T, 465 A, 579 G, 582 G, 591 G, 648 G, 678 GC
This species has been reared from Carathis septentrionalis (Erebidae) feeding on mature leaves of Ocotea cernua (Lauraceae) and Pachydota saduca (Erebidae) feeding on several species of Ocotea and Nectandra (Lauraceae). Host caterpillars of type specimens were collected throughout the year, except between March and May.
Holotype ♀: DHJPAR0036333 (ASHYD1524-09), Costa Rica, Área de Conservación Guanacaste, Sector Rincon Rain Forest, Estacion Llanura, GPS: 10.93332, -85.25331, 135 m elevation, Keiner Aragon coll., reared from Pachydota saduca 09-SRNP-44900, host collected 4 July 2009, wasp eclosed 8 September 2009, (
Hemichoma johnkingsolveri is named in honor of John Kingsolver (RIP) of the USDA Systematic Entomology Laboratory at the Smithsonian Institution, a master taxonomist of Bruchinae (Chrysomelidae) and long-time supporter of ACG biodiversity inventory.
Ichneumonoid taxonomists have remained in a taxonomic paradigm that was created for a well-known and largely extra-tropical fauna and flora. For example, there is great utility in a morphological key to the 30 species of butterflies that occur in a suburban backyard in eastern North America; however a key to the 100+ species of 1–4 cm long amber-colored nocturnal highly host-specific species of Enicospilus parasitic wasps (Ichneumonidae: Ophioninae) that occur within 3 km of the Administration Area of Área de Conservación Guanacaste is much less useful because, a) they mostly look the same, b) 90% are undescribed, and c) knowing the species name would not give you much additional information, i.e., life history, geographic range, or phenology. Now there is an alternative to morphological keys and complex prose descriptions (
With online public databases such as BOLD accumulating hundreds of thousands of specimen and species based barcodes (
We recognize that DNA barcodes may fail to delimit all species, or all specimens of all species, just as there are no morphological characters capable of unfailingly separating species. There are drawbacks to using a portion of the single gene COI as a barcode, including potential confusion with nuclear mitochondrial paralogs, Wolbachia mediated introgression, hybridization, and incomplete lineage sorting (
We demonstrate a novel approach to species descriptions for hyperdiverse, underdescribed taxa, such as those within Ichneumonoidea. The descriptions consist of a high-quality lateral habitus photograph, latitude and longitude coordinates, and the diagnostic characteristics of the COI barcode region, along with details of the holotype as required by the International Code of Zoological Nomenclature (
We gratefully acknowledge the unflagging support of the team of ACG parataxonomists (