DNA barcodes, like traditional sources of taxonomic information, are potentially powerful heuristics in the identification of described species but require mindful analytical interpretation. The role of DNA barcoding in generating hypotheses of new taxa in need of formal taxonomic treatment is discussed, and it is emphasized that the recursive process of character evaluation is both necessary and best served by understanding the empirical mechanics of the discovery process. These undertakings carry enormous ramifications not only for the translation of DNA sequence data into taxonomic information but also for our comprehension of the magnitude of species diversity and its disappearance. This paper examines the potential strengths and pitfalls of integrating DNA sequence data, specifically in the form of DNA barcodes as they are currently generated and analyzed, with taxonomic practice.
Three new genera are described: Michener (Proteropinae), Bioalfa (Rogadinae), and Hermosomastax (Rogadinae). Keys are given for the New World genera of the following braconid subfamilies: Agathidinae, Braconinae, Cheloninae, Homolobinae, Hormiinae, Ichneutinae, Macrocentrinae, Orgilinae, Proteropinae, Rhysipolinae, and Rogadinae. In these subfamilies 416 species are described or redescribed. Most of the species have been reared and all but 13 are new to science. A consensus sequence of the COI barcodes possessed by each species is employed to diagnose the species, and this approach is justified in the introduction. Most descriptions consist of a lateral or dorsal image of the holotype, a diagnostic COI consensus barcode, the Barcode Index Number (BIN) code with a link to the Barcode of Life Database (BOLD), and the holotype specimen information required by the International Code of Zoological Nomenclature. The following species are treated and those lacking authorship are newly described here with authorship attributable to Sharkey except for the new species of Macrocentrinae which are by Sharkey & van Achterberg: AGATHIDINAE: Aerophilus paulmarshi, Mesocoelus davidsmithi, Neothlipsis bobkulai, Plesiocoelus vanachterbergi, Pneumagathis erythrogastra (Cameron, 1905), Therophilus bobwhartoni, T. donaldquickei, T. gracewoodae, T. maetoi, T. montywoodi, T. penteadodiasae, Zacremnops brianbrowni, Z. coatlicue Sharkey, 1990, Zacremnops cressoni (Cameron, 1887), Z. ekchuah Sharkey, 1990, Z. josefernandezi, Zelomorpha sarahmeierottoae. BRACONINAE: Bracon alejandromarini, B. alejandromasisi, B. alexamasisae, B. andresmarini, B. andrewwalshi, B. anniapicadoae, B. anniemoriceae, B. barryhammeli, B. bernardoespinozai, B. carlossanabriai, B. chanchini, B. christophervallei, B. erasmocoronadoi, B. eugeniephillipsae, B. federicomatarritai, B. frankjoycei, B. gerardovegai, B. germanvegai, B. isidrochaconi, B. jimlewisi, B. josejaramilloi, B. juanjoseoviedoi, B. juliodiazi, B. luzmariaromeroae, B. manuelzumbadoi, B. marialuisariasae, B. mariamartachavarriae, B. mariorivasi, B. melissaespinozae, B. nelsonzamorai, B. nicklaphami, B. ninamasisae, B. oliverwalshi, B. paulamarinae, B. rafamoralesi, B. robertofernandezi, B. rogerblancoi, B. ronaldzunigai, B. sigifredomarini, B. tihisiaboshartae, B. wilberthbrizuelai, Digonogastra montylloydi, D. montywoodi, D. motohasegawai, D. natwheelwrighti, D. nickgrishini. CHELONINAE: Adelius adrianguadamuzi, A. gauldi Shimbori & Shaw, 2019, A. janzeni Shimbori & Shaw, 2019, Ascogaster gloriasihezarae, A. grettelvegae, A. guillermopereirai, A. gustavoecheverrii, A. katyvandusenae, A. luisdiegogomezi, Chelonus alejandrozaldivari, C. gustavogutierrezi, C. gustavoinduni, C. harryramirezi, C. hartmanguidoi, C. hazelcambroneroae, C. iangauldi, C. isidrochaconi, C. janecheverriae, C. jeffmilleri, C. jennyphillipsae, C. jeremydewaardi, C. jessiehillae, C. jesusugaldei, C. jimlewisi, C. jimmilleri, C. jimwhitfieldi, C. johanvalerioi, C. johnburnsi, C. johnnoyesi, C. jorgebaltodanoi, C. jorgehernandezi, C. josealfredohernandezi, C. josefernandeztrianai, C. josehernandezcortesi, C. josemanuelperezi, C. josephinerodriguezae, C. juanmatai, C. junkoshimurae, C. kateperezae, C. luciariosae, C. luzmariaromeroae, C. manuelpereirai, C. manuelzumbadoi, C. marianopereirai, C. maribellealvarezae, C. markmetzi, C. markshawi, C. martajimenezae, C. mayrabonillae, C. meganmiltonae, C. melaniamunozae, C. michaelstroudi, C. michellevanderbankae, C. mingfangi, C. minorcarmonai, C. monikaspringerae, C. moniquegilbertae, C. motohasegawai, C. nataliaivanovae, C. nelsonzamorai, C. normwoodleyi, C. osvaldoespinozai, C. pamelacastilloae, C. paulgoldsteini, C. paulhansoni, C. paulheberti, C. petronariosae, C. ramyamanjunathae, C. randallgarciai, C. rebeccakittelae, C. robertoespinozai, C. robertofernandezi, C. rocioecheverriae, C. rodrigogamezi, C. ronaldzunigai, C. rosibelelizondoae, C. rostermoragai, C. ruthfrancoae, C. scottmilleri, C. scottshawi, C. sergioriosi, C. sigifredomarini, C. stevearonsoni, C. stevestroudi, C. sujeevanratnasinghami, C. sureshnaiki, C. torbjornekremi, C. yeimycedenoae, Leptodrepana alexisae, L. erasmocoronadoi, L. felipechavarriai, L. freddyquesadai, L. gilbertfuentesi, L. manuelriosi, Phanerotoma almasolisae, P. alvaroherrerai, P. anacordobae, P. anamariamongeae, P. andydeansi, P. angelagonzalezae, P. angelsolisi, P. barryhammeli, P. bernardoespinozai, P. calixtomoragai, P. carolinacanoae, P. christerhanssoni, P. christhompsoni, P. davesmithi, P. davidduthiei, P. dirksteinkei, P. donquickei, P. duniagarciae, P. duvalierbricenoi, P. eddysanchezi, P. eldarayae, P. eliethcantillanoae, P. jenopappi, Pseudophanerotoma alanflemingi, Ps. albanjimenezi, Ps. alejandromarini, Ps. alexsmithi, Ps. allisonbrownae, Ps. bobrobbinsi. HOMOLOBINAE: Exasticolus jennyphillipsae, E. randallgarciai, E. robertofernandezi, E. sigifredomarini, E. tomlewinsoni. HORMIINAE: Hormius anamariamongeae, H. angelsolisi, H. anniapicadoae, H. arthurchapmani, H. barryhammeli, H. carmenretanae, H. carloswalkeri, H. cesarsuarezi, H. danbrooksi, H. eddysanchezi, H. erikframstadi, H. georgedavisi, H. grettelvegae, H. gustavoinduni, H. hartmanguidoi, H. hectoraritai, H. hesiquiobenitezi, H. irenecanasae, H. isidrochaconi, H. jaygallegosi, H. jimbeachi, H. jimlewisi, H. joelcracrafti, H. johanvalerioi, H. johnburleyi, H. joncoddingtoni, H. jorgecarvajali, H. juanmatai, H. manuelzumbadoi, H. mercedesfosterae, H. modonnellyae, H. nelsonzamorai, H. pamelacastilloae, H. raycypessi, H. ritacolwellae, H. robcolwelli, H. rogerblancosegurai, H. ronaldzunigai, H. russchapmani, H. virginiaferrisae, H. warrenbrighami, H. willsflowersi. ICHNEUTINAE: Oligoneurus kriskrishtalkai, O. jorgejimenezi, Paroligoneurus elainehoaglandae, P. julianhumphriesi, P. mikeiviei. MACROCENTRINAE: Austrozele jorgecampabadali, A. jorgesoberoni, Dolichozele gravitarsis (Muesebeck, 1938), D. josefernandeztrianai, D. josephinerodriguezae, Hymenochaonia kalevikulli, H. kateperezae, H. katherinebaillieae, H. katherineellisonae, H. katyvandusenae, H. kazumifukunagae, H. keithlangdoni, H. keithwillmotti, H. kenjinishidai, H. kimberleysheldonae, H. krisnorvigae, H. lilianamadrigalae, H. lizlangleyae, Macrocentrus fredsingeri, M. geoffbarnardi, M. gregburtoni, M. gretchendailyae, M. grettelvegae, M. gustavogutierrezi, M. hannahjamesae, M. harisridhari, M. hillaryrosnerae, M. hiroshikidonoi, M. iangauldi, M. jennyphillipsae, M. jesseausubeli, M. jessemaysharkae, M. jimwhitfieldi, M. johnbrowni, M. johnburnsi, M. jonathanfranzeni, M. jonathanrosenbergi, M. jorgebaltodanoi, M. lucianocapelli. ORGILINAE: Orgilus amyrossmanae, O. carrolyoonae, O. christhompsoni, O. christinemcmahonae, O. dianalipscombae, O. ebbenielsoni, O. elizabethpennisiae, O. evertlindquisti, O. genestoermeri, O. jamesriegeri, O. jeanmillerae, O. jeffmilleri, O. jerrypowelli, O. jimtiedjei, O. johnlundbergi, O. johnpipolyi, O. jorgellorentei, O. larryspearsi, O. marlinricei, O. mellissaespinozae, O. mikesmithi, O. normplatnicki, O. peterrauchi, O. richardprimacki, O. sandraberriosae, O. sarahmirandae, O. scottmilleri, O. scottmorii, Stantonia billalleni, S. brookejarvisae, S. donwilsoni, S. erikabjorstromae, S. garywolfi, S. henrikekmani, S. luismirandai, S. miriamzunzae, S. quentinwheeleri, S. robinkazmierae, S. ruthtifferae. PROTEROPINAE: Hebichneutes tricolor Sharkey & Wharton, 1994, Proterops iangauldi, P. vickifunkae, Michener charlesi. RHYSIPOLINAE: Pseudorhysipolis luisfonsecai, P. mailyngonzalezaeRhysipolis julioquirosi. ROGADINAE: Aleiodes adrianaradulovae, A. adrianforsythi, A. agnespeelleae, A. alaneaglei, A. alanflemingi, A. alanhalevii, A. alejandromasisi, A. alessandracallejae, A. alexsmithi, A. alfonsopescadori, A. alisundermieri, A. almasolisae, A. alvarougaldei, A. alvaroumanai, A. angelsolisi, A. annhowdenae, A. bobandersoni, A. carolinagodoyae, A. charlieobrieni, A. davefurthi, A. donwhiteheadi, A. doylemckeyi, A. frankhovorei, A. henryhowdeni, A. inga Shimbori & Shaw, 2020, A. johnchemsaki, A. johnkingsolveri, A. gonodontovorus Shimbori & Shaw, 2020, A. manuelzumbadoi, A. mayrabonillae, A. michelledsouzae, A. mikeiviei, A. normwoodleyi, A. pammitchellae, A. pauljohnsoni, A. rosewarnerae, A. steveashei, A. terryerwini, A. willsflowersi, Bioalfa pedroleoni, B. alvarougaldei, B. rodrigogamezi, Choreborogas andydeansi, C. eladiocastroi, C. felipechavarriai, C. frankjoycei, Clinocentrus andywarreni, Cl. angelsolisi, Cystomastax alexhausmanni, Cy. angelagonzalezae, Cy. ayaigarashiae, Hermosomastax clavifemorus Quicke sp. nov., Heterogamus donstonei, Pseudoyelicones bernsweeneyi, Stiropius bencrairi, S. berndkerni, S. edgargutierrezi, S. edwilsoni, S. ehakernae, Triraphis billfreelandi, T. billmclarneyi, T. billripplei, T. bobandersoni, T. bobrobbinsi, T. bradzlotnicki, T. brianbrowni, T. brianlaueri, T. briannestjacquesae, T. camilocamargoi, T. carlosherrerai, T. carolinepalmerae, T. charlesmorrisi, T. chigiybinellae, T. christerhanssoni, T. christhompsoni, T. conniebarlowae, T. craigsimonsi, T. defectus Valerio, 2015, T. danielhubi, T. davidduthiei, T. davidwahli, T. federicomatarritai, T. ferrisjabri, T. mariobozai, T. martindohrni, T. matssegnestami, T. mehrdadhajibabaei, T. ollieflinti, T. tildalauerae, Yelicones dirksteinkei, Y. markmetzi, Y. monserrathvargasae, Y. tricolor Quicke, 1996. Y. woldai Quicke, 1996. The following new combinations are proposed: Neothlipsis smithi (Ashmead), new combination for Microdus smithi Ashmead, 1894; Neothlipsis pygmaeus (Enderlein), new combination for Microdus pygmaeus Enderlein, 1920; Neothlipsis unicinctus (Ashmead), new combination for Microdus unicinctus Ashmead, 1894; Therophilus anomalus (Bortoni and Penteado-Dias) new combination for Plesiocoelus anomalus Bortoni and Penteado-Dias, 2015; Aerophilus areolatus (Bortoni and Penteado-Dias) new combination for Plesiocoelus areolatus Bortoni and Penteado-Dias, 2015; Pneumagathis erythrogastra (Cameron) new combination for Agathis erythrogastra Cameron, 1905. Dolichozele citreitarsis (Enderlein), new combination for Paniscozele citreitarsis Enderlein, 1920. Dolichozele fuscivertex (Enderlein) new combination for Paniscozele fuscivertex Enderlein, 1920. Finally, Bassus brooksi Sharkey, 1998 is synonymized with Agathis erythrogastra Cameron, 1905; Paniscozele griseipes Enderlein, 1920 is synonymized with Dolichozele koebelei Viereck, 1911; Paniscozele carinifrons Enderlein, 1920 is synonymized with Dolichozele fuscivertex (Enderlein, 1920); and Paniscozele nigricauda Enderlein,1920 is synonymized with Dolichozele quaestor (Fabricius, 1804). (originally described as Ophion quaestor Fabricius, 1804).
Interpretations and analytical practices surrounding DNA barcoding are examined using a compilation of 3,756 papers (as of December 31, 2018) with "DNA Barcode" in the abstract published since 2004. By examining the rise of DNA barcoding in natural history and biodiversity science over this period, we hope to detect the extent to which its purposes, premises, rationale and application have evolved. The number of studies involving identification, taxonomic decisions and the discovery of cryptic species has grown rapidly and appears to have driven much of the publication activity of DNA barcode studies overall. Forensic studies and papers on biological conservation involving DNA barcodes have loosely tracked the ensemble number of studies but appear to have risen sharply in 2017. Although analytical paradigms have diversified, particularly following the growing availability of tools in BoLD, neighbor-joining and graphic (tree-based) criteria for species delimitation remain preeminent. We conclude that the practices and paradigms of DNA barcoding data are likely to persist and, in groups such as Lepidoptera, remain a widely used tool in taxonomic science.
Conservation genetics has expanded its purview such that molecular techniques are now used routinely to prioritize populations for listing and protection and infer their historical relationships in addition to addressing more traditional questions of heterozygosity and inbreeding depression. Failure to specify whether molecular data are being used for diagnosis‐related questions or for population viability questions, however, can lead either to misinterpretation of character data as adaptive information or to misinterpretation of frequency or distance data as diagnostic or historical information. Each of these misinterpretations will confound conservation programs. The character‐based approach to delimiting phylogenetic species is both operationally and logically superior to “diagnostic” methods that involve distance‐ or frequency‐based routines, which are unstable over time. Tree‐based criteria for the diagnosis of conservation “units” are also inappropriate because they can depend on patterns inferred without reference to diagnostic characters. Intraspecific studies, conservation‐related or otherwise, that adopt terminology and methods designed to infer nested hierarchic relationships confuse diagnosis with historical inferences by treating diagnoses as outcomes rather than as precursors to phylogeny reconstruction. A character‐based diagnostic approach recognizes the analytical dichotomy between species hierarchies and population statistics and provides a framework for the understanding of each. No species concept, however, should be viewed as an absolute criterion for protecting populations, but as part of a framework from within which identification of protection and management goals can be achieved effectively and defensibly.
Museum specimens from the late 19th and early 20th centuries were surveyed for the single nucleotide polymorphism identified previously and used to diagnose populations of the federally threatened Northeastern Beach Tiger Beetle Cicindela d. dorsalis (Coleoptera: Carabidae). Widespread polymorphism was revealed throughout the historical range of this species, suggesting a relatively recent anthropogenic character fixation event associated with the extinction and fragmentation of populations. Implications for the phylogenetic species criterion and for the reintroduction of individuals to formerly occupied sites are discussed.
Several increasingly popular paradigms in conservation remove organismal information and life‐history requirements from management planning with claims that species‐specific information is not necessary to the understanding and management of “ecosystem function” and may therefore be discarded. Although several authors have called attention to the fact that ecosystem management has not yet been articulated sufficiently to comprise an adequate paradigm for wildlife protection, there has been a series of suggestions paralleling ecosystem management’s popularity that the perceived or imagined emergent properties of communities should be at the root of conservation planning. Such reductions most commonly take the form of abstracted species diversity measures that may be irrelevant or misleading with respect to site‐specific planning and to the monitoring of specific management treatments. Following earlier examinations of ecosystem management, I emphasize that several of its apparent outgrowths may be too vague to inform specific recommendations, that the historical mechanics of “ecosystem processes” are essentially unknowable, and that anecdotal definitions of ecosystems allow one to justify virtually any protocol as management. The primacy of process‐dependent, landscape‐functional considerations in conservation planning is specious in the absence of species‐ and population‐specific information, which should form the foundation for understanding such processes in the first place. Only by viewing nature in terms of the relationships between processes and organisms, rather than in terms of emergent properties of organismal assemblages or abiotic factors divorced from organismal data, can conservation plans claim to protect biological elements.
Molecular identification is increasingly used to speed up biodiversity surveys and laboratory experiments. However, many groups of organisms cannot be reliably identified using standard databases such as GenBank or BOLD due to lack of sequenced voucher specimens identified by experts. Sometimes a large number of sequences are available, but with too many errors to allow identification. Here we address this problem for parasitoids of Drosophila by introducing a curated open-access molecular reference database, DROP (Drosophila parasitoids). Identifying Drosophila parasitoids is specimens are identified by taxonomists and vetted through direct comparison with primary type material. To initiate DROP, we curated 154 laboratory strains, 853 vouchers, 545 DNA sequences, 16 genomes, 11 transcriptomes, and 6 proteomes drawn from a total of 183 operational taxonomic units (OTUs): 113 described Drosophila parasitoid species and 70 provisional species. We found species richness of Drosophila parasitoids to be acutely underestimated and provide an updated taxonomic catalogue for the community. DROP offers accurate molecular identification and improves crossreferencing between individual studies that we hope will catalyze research on this diverse and fascinating model system. Our effort should also serve as an example for researchers facing similar molecular identification problems in other groups of organisms.
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