Imagine if we could compute across phenotype data as easily as genomic data; this article calls for efforts to realize this vision and discusses the potential benefits.
Wolbachia is a genus of bacterial endosymbionts that impacts the breeding systems of their hosts. Wolbachia can confuse the patterns of mitochondrial variation, including DNA barcodes, because it influences the pathways through which mitochondria are inherited. We examined the extent to which these endosymbionts are detected in routine DNA barcoding, assessed their impact upon the insect sequence divergence and identification accuracy, and considered the variation present in Wolbachia COI. Using both standard PCR assays (Wolbachia surface coding protein – wsp), and bacterial COI fragments we found evidence of Wolbachia in insect total genomic extracts created for DNA barcoding library construction. When >2 million insect COI trace files were examined on the Barcode of Life Datasystem (BOLD) Wolbachia COI was present in 0.16% of the cases. It is possible to generate Wolbachia COI using standard insect primers; however, that amplicon was never confused with the COI of the host. Wolbachia alleles recovered were predominantly Supergroup A and were broadly distributed geographically and phylogenetically. We conclude that the presence of the Wolbachia DNA in total genomic extracts made from insects is unlikely to compromise the accuracy of the DNA barcode library; in fact, the ability to query this DNA library (the database and the extracts) for endosymbionts is one of the ancillary benefits of such a large scale endeavor – for which we provide several examples. It is our conclusion that regular assays for Wolbachia presence and type can, and should, be adopted by large scale insect barcoding initiatives. While COI is one of the five multi-locus sequence typing (MLST) genes used for categorizing Wolbachia, there is limited overlap with the eukaryotic DNA barcode region.
Frequently, the diversity of umbrella taxa is invoked to predict patterns of other, less well-known, life. However, the utility of this strategy has been questioned. We tested whether a phylogenetic diversity (PD) analysis of CO1 DNA barcodes could act as a proxy for standard methods of determining sampling efficiency within and between sites, namely that an accumulation curve of barcode diversity would be similar to curves generated using morphology or nuclear genetic markers. Using taxa at the forefront of the taxonomic impediment -parasitoid wasps (Ichneumonidae, Braconidae, Cynipidae and Diapriidae), contrasted with a taxon expected to be of low diversity (Formicidae) from an area where total diversity is expected to be low (Churchill, Manitoba), we found that barcode accumulation curves based on PD were significantly different in both slope and scale from curves generated using names based on morphological data, while curves generated using nuclear genetic data were only different in scale. We conclude that these differences clearly identify the taxonomic impediment within the strictly morphological alpha-taxonomy of these hyperdiverse insects. The absence of an asymptote within the barcode PD trend of parasitoid wasps reflects the as yet incomplete sampling of the site (and more accurately its total diversity), while the morphological analysis asymptote represents a collision with the taxonomic impediment rather than complete sampling. We conclude that a PD analysis of standardized DNA barcodes can be a transparent and reproducible triage tool for the management and conservation of species and spaces.
More than half a million specimens of wild-caught Lepidoptera caterpillars have been reared for their parasitoids, identified, and DNA barcoded over a period of 34 years (and ongoing) from Area de Conservación de Guanacaste (ACG), northwestern Costa Rica. This provides the world’s best location-based dataset for studying the taxonomy and host relationships of caterpillar parasitoids. Among Hymenoptera, Microgastrinae (Braconidae) is the most diverse and commonly encountered parasitoid subfamily, with many hundreds of species delineated to date, almost all undescribed. Here, we reassess the limits of the genus Apanteles sensu stricto, describe 186 new species from 3,200+ parasitized caterpillars of hundreds of ACG Lepidoptera species, and provide keys to all 205 described Apanteles from Mesoamerica – including 19 previously described species in addition to the new species. The Mesoamerican Apanteles are assigned to 32 species-groups, all but two of which are newly defined. Taxonomic keys are presented in two formats: traditional dichotomous print versions and links to electronic interactive versions (software Lucid 3.5). Numerous illustrations, computer-generated descriptions, distributional information, wasp biology, and DNA barcodes (where available) are presented for every species. All morphological terms are detailed and linked to the Hymenoptera Anatomy Ontology website. DNA barcodes (a standard fragment of the cytochrome c oxidase I (COI) mitochondrial gene), information on wasp biology (host records, solitary/gregariousness of wasp larvae), ratios of morphological features, and wasp microecological distributions were used to help clarify boundaries between morphologically cryptic species within species-complexes. Because of the high accuracy of host identification for about 80% of the wasp species studied, it was possible to analyze host relationships at a regional level. The ACG species of Apanteles attack mainly species of Hesperiidae, Elachistidae and Crambidae (Lepidoptera). About 90% of the wasp species with known host records seem to be monophagous or oligophagous at some level, parasitizing just one host family and commonly, just one species of caterpillar. Only 15 species (9%) parasitize species in more than one family, and some of these cases are likely to be found to be species complexes. We have used several information sources and techniques (traditional taxonomy, molecular, software-based, biology, and geography) to accelerate the process of finding and describing these new species in a hyperdiverse group such as Apanteles.The following new taxonomic and nomenclatural acts are proposed. Four species previously considered to be Apanteles are transferred to other microgastrine genera: Dolichogenidea hedyleptae (Muesebeck, 1958), comb. n., Dolichogenidea politiventris (Muesebeck, 1958), comb. n., Rhygoplitis sanctivincenti (Ashmead, 1900), comb. n., and Illidops scutellaris (Muesebeck, 1921), comb. rev. One European species that is a secondary homonym to a Mesoamerican species is removed from Apan...
Climate change will have profound and unanticipated effects on species distributions. The pace and nature of this change is largely unstudied, especially for the most diverse elements of terrestrial communities – the arthropods – here we have only limited knowledge concerning the taxonomy and the ecology of these groups. Because Arctic ecosystems have already experienced significant increases in temperature over the past half century, shifts in community structure may already be in progress. Here we utilise collections of a particularly hyperdiverse insect group – parasitoid wasps (Hymenoptera; Braconidae; Microgastrinae) – at Churchill, Manitoba, Canada in the early and mid-twentieth century to compare the composition of the contemporary community to that present 50–70 years ago. Morphological and DNA barcoding results revealed the presence of 79 species of microgastrine wasps in collections from Churchill, but we estimate that 20% of the local fauna awaits detection. Species composition and diversity between the two time periods differ significantly; species that were most common in historic collections were not found in contemporary collections and vice versa. Using barcodes we compared these collections to others from across North America; contemporary Churchill species are most affiliated with more south-western collections, while historic collections were more affiliated with eastern collections. The past five decades has clearly seen a dramatic change of species composition within the area studied coincident with rising temperature.
A checklist of world species of Microgastrinae parasitoid wasps (Hymenoptera: Braconidae) is provided. A total of 81 genera and 2,999 extant species are recognized as valid, including 36 nominal species that are currently considered as species inquirendae. Two genera are synonymized under Apanteles. Nine lectotypes are designated. A total of 318 new combinations, three new replacement names, three species name amendments, and seven species status revised are proposed. Additionally, three species names are treated as nomina dubia, and 52 species names are considered as unavailable names (including 14 as nomina nuda). A total of three extinct genera and 12 extinct species are also listed. Unlike in many previous treatments of the subfamily, tribal concepts are judged to be inadequate, so genera are listed alphabetically. Brief diagnoses of all Microgastrinae genera, as understood in this paper, are presented. Illustrations of all extant genera (at least one species per genus, usually more) are included to showcase morphological diversity. Primary types of Microgastrinae are deposited in 108 institutions worldwide, although 76% are concentrated in 17 collections. Localities of primary types, in 138 countries, are reported. Recorded species distributions are listed by biogeographical region and by country. Microgastrine wasps are recorded from all continents except Antarctica; specimens can be found in all major terrestrial ecosystems, from 82°N to 55°S, and from sea level up to at least 4,500 m a.s.l. The Oriental (46) and Neotropical (43) regions have the largest number of genera recorded, whereas the Palaearctic region (28) is the least diverse. Currently, the highest species richness is in the Palearctic region (827), due to more historical study there, followed by the Neotropical (768) and Oriental (752) regions, which are expected to be the most species rich. Based on ratios of Lepidoptera and Microgastrinae species from several areas, the actual world diversity of Microgastrinae is expected to be between 30,000–50,000 species; although these ratios were mostly based on data from temperate areas and thus must be treated with caution, the single tropical area included had a similar ratio to the temperate ones. Almost 45,000 specimens of Microgastrinae from 67 different genera (83% of microgastrine genera) have complete or partial DNA barcode sequences deposited in the Barcode of Life Data System; the DNA barcodes represent 3,545 putative species or Barcode Index Numbers (BINs), as estimated from the molecular data. Information on the number of sequences and BINs per genus are detailed in the checklist. Microgastrinae hosts are here considered to be restricted to Eulepidoptera, i.e., most of the Lepidoptera except for the four most basal superfamilies (Micropterigoidea, Eriocranioidea, Hepialoidea and Nepticuloidea), with all previous literature records of other insect orders and those primitive Lepidoptera lineages being considered incorrect. The following nomenclatural acts are proposed: 1) Two genera are synonymyzed under Apanteles: Cecidobracon Kieffer & Jörgensen, 1910, new synonym and Holcapanteles Cameron, 1905, new synonym; 2) Nine lectotype designations are made for Alphomelon disputabile (Ashmead, 1900), Alphomelon nigriceps (Ashmead, 1900), Cotesia salebrosa (Marshall, 1885), Diolcogaster xanthaspis (Ashmead, 1900), Dolichogenidea ononidis (Marshall, 1889), Glyptapanteles acraeae (Wilkinson, 1932), Glyptapanteles guyanensis (Cameron, 1911), Glyptapanteles militaris (Walsh, 1861), and Pseudapanteles annulicornis Ashmead, 1900; 3) Three new replacement names are a) Diolcogaster aurangabadensis Fernandez-Triana, replacing Diolcogaster indicus (Rao & Chalikwar, 1970) [nec Diolcogaster indicus (Wilkinson, 1927)], b) Dolichogenidea incystatae Fernandez-Triana, replacing Dolichogenidea lobesia Liu & Chen, 2019 [nec Dolichogenidea lobesia Fagan-Jeffries & Austin, 2019], and c) Microplitis vitobiasi Fernandez-Triana, replacing Microplitis variicolor Tobias, 1964 [nec Microplitis varicolor Viereck, 1917]; 4) Three names amended are Apanteles irenecarrilloae Fernandez-Triana, 2014, Cotesia ayerzai (Brèthes, 1920), and Cotesia riverai (Porter, 1916); 5) Seven species have their status revised: Cotesia arctica (Thomson, 1895), Cotesia okamotoi (Watanabe, 1921), Cotesia ukrainica (Tobias, 1986), Dolichogenidea appellator (Telenga, 1949), Dolichogenidea murinanae (Capek & Zwölfer, 1957), Hypomicrogaster acarnas Nixon, 1965, and Nyereria nigricoxis (Wilkinson, 1932); 6) New combinations are given for 318 species: Alloplitis congensis, Alloplitis detractus, Apanteles asphondyliae, Apanteles braziliensis, Apanteles sulciscutis, Choeras aper, Choeras apollion, Choeras daphne, Choeras fomes, Choeras gerontius, Choeras helle, Choeras irates, Choeras libanius, Choeras longiterebrus, Choeras loretta, Choeras recusans, Choeras sordidus, Choeras stenoterga, Choeras superbus, Choeras sylleptae, Choeras vacillatrix, Choeras vacillatropsis, Choeras venilia, Cotesia asavari, Cotesia bactriana, Cotesia bambeytripla, Cotesia berberidis, Cotesia bhairavi, Cotesia biezankoi, Cotesia bifida, Cotesia caligophagus, Cotesia cheesmanae, Cotesia compressithorax, Cotesia delphinensis, Cotesia effrena, Cotesia euphobetri, Cotesia elaeodes, Cotesia endii, Cotesia euthaliae, Cotesia exelastisae, Cotesia hiberniae, Cotesia hyperion, Cotesia hypopygialis, Cotesia hypsipylae, Cotesia jujubae, Cotesia lesbiae, Cotesia levigaster, Cotesia lizeri, Cotesia malevola, Cotesia malshri, Cotesia menezesi, Cotesia muzaffarensis, Cotesia neptisis, Cotesia nycteus, Cotesia oeceticola, Cotesia oppidicola, Cotesia opsiphanis, Cotesia pachkuriae, Cotesia paludicolae, Cotesia parbhanii, Cotesia parvicornis, Cotesia pratapae, Cotesia prozorovi, Cotesia pterophoriphagus, Cotesia radiarytensis, Cotesia rangii, Cotesia riverai, Cotesia ruficoxis, Cotesia senegalensis, Cotesia seyali, Cotesia sphenarchi, Cotesia sphingivora, Cotesia transuta, Cotesia turkestanica, Diolcogaster abengouroui, Diolcogaster agama, Diolcogaster ambositrensis, Diolcogaster anandra, Diolcogaster annulata, Diolcogaster bambeyi, Diolcogaster bicolorina, Diolcogaster cariniger, Diolcogaster cincticornis, Diolcogaster cingulata, Diolcogaster coronata, Diolcogaster coxalis, Diolcogaster dipika, Diolcogaster earina, Diolcogaster epectina, Diolcogaster epectinopsis, Diolcogaster grangeri, Diolcogaster heterocera, Diolcogaster homocera, Diolcogaster indica, Diolcogaster insularis, Diolcogaster kivuana, Diolcogaster mediosulcata, Diolcogaster megaulax, Diolcogaster neglecta, Diolcogaster nigromacula, Diolcogaster palpicolor, Diolcogaster persimilis, Diolcogaster plecopterae, Diolcogaster plutocongoensis, Diolcogaster psilocnema, Diolcogaster rufithorax, Diolcogaster semirufa, Diolcogaster seyrigi, Diolcogaster subtorquata, Diolcogaster sulcata, Diolcogaster torquatiger, Diolcogaster tristiculus, Diolcogaster turneri, Diolcogaster vulcana, Diolcogaster wittei, Distatrix anthedon, Distatrix cerales, Distatrix cuspidalis, Distatrix euproctidis, Distatrix flava, Distatrix geometrivora, Distatrix maia, Distatrix tookei, Distatrix termina, Distatrix simulissima, Dolichogenidea agamedes, Dolichogenidea aluella, Dolichogenidea argiope, Dolichogenidea atreus, Dolichogenidea bakeri, Dolichogenidea basiflava, Dolichogenidea bersa, Dolichogenidea biplagae, Dolichogenidea bisulcata, Dolichogenidea catonix, Dolichogenidea chrysis, Dolichogenidea coffea, Dolichogenidea coretas, Dolichogenidea cyane, Dolichogenidea diaphantus, Dolichogenidea diparopsidis, Dolichogenidea dryas, Dolichogenidea earterus, Dolichogenidea ensiger, Dolichogenidea eros, Dolichogenidea evadne, Dolichogenidea falcator, Dolichogenidea gelechiidivoris, Dolichogenidea gobica, Dolichogenidea hyalinis, Dolichogenidea iriarte, Dolichogenidea lakhaensis, Dolichogenidea lampe, Dolichogenidea laspeyresiella, Dolichogenidea latistigma, Dolichogenidea lebene, Dolichogenidea lucidinervis, Dolichogenidea malacosomae, Dolichogenidea maro, Dolichogenidea mendosae, Dolichogenidea monticola, Dolichogenidea nigra, Dolichogenidea olivierellae, Dolichogenidea parallelis, Dolichogenidea pelopea, Dolichogenidea pelops, Dolichogenidea phaenna, Dolichogenidea pisenor, Dolichogenidea roepkei, Dolichogenidea scabra, Dolichogenidea statius, Dolichogenidea stenotelas, Dolichogenidea striata, Dolichogenidea wittei, Exoryza asotae, Exoryza belippicola, Exoryza hylas, Exoryza megagaster, Exoryza oryzae, Glyptapanteles aggestus, Glyptapanteles agynus, Glyptapanteles aithos, Glyptapanteles amenophis, Glyptapanteles antarctiae, Glyptapanteles anubis, Glyptapanteles arginae, Glyptapanteles argus, Glyptapanteles atylana, Glyptapanteles badgleyi, Glyptapanteles bataviensis, Glyptapanteles bistonis, Glyptapanteles borocerae, Glyptapanteles cacao, Glyptapanteles cadei, Glyptapanteles cinyras, Glyptapanteles eryphanidis, Glyptapanteles euproctisiphagus, Glyptapanteles eutelus, Glyptapanteles fabiae, Glyptapanteles fulvigaster, Glyptapanteles fuscinervis, Glyptapanteles gahinga, Glyptapanteles globatus, Glyptapanteles glyphodes, Glyptapanteles guierae, Glyptapanteles horus, Glyptapanteles intricatus, Glyptapanteles lamprosemae, Glyptapanteles lefevrei, Glyptapanteles leucotretae, Glyptapanteles lissopleurus, Glyptapanteles madecassus, Glyptapanteles marquesi, Glyptapanteles melanotus, Glyptapanteles melissus, Glyptapanteles merope, Glyptapanteles naromae, Glyptapanteles nepitae, Glyptapanteles nigrescens, Glyptapanteles ninus, Glyptapanteles nkuli, Glyptapanteles parasundanus, Glyptapanteles penelope, Glyptapanteles penthocratus, Glyptapanteles philippinensis, Glyptapanteles philocampus, Glyptapanteles phoebe, Glyptapanteles phytometraduplus, Glyptapanteles propylae, Glyptapanteles puera, Glyptapanteles seydeli, Glyptapanteles siderion, Glyptapanteles simus, Glyptapanteles speciosissimus, Glyptapanteles spilosomae, Glyptapanteles subpunctatus, Glyptapanteles thespis, Glyptapanteles thoseae, Glyptapanteles venustus, Glyptapanteles wilkinsoni, Hypomicrogaster samarshalli, Iconella cajani, Iconella detrectans, Iconella jason, Iconella lynceus, Iconella pyrene, Iconella tedanius, Illidops azamgarhensis, Illidops lamprosemae, Illidops trabea, Keylimepie striatus, Microplitis adisurae, Microplitis mexicanus, Neoclarkinella ariadne, Neoclarkinella curvinervus, Neoclarkinella sundana, Nyereria ituriensis, Nyereria nioro, Nyereria proagynus, Nyereria taoi, Nyereria vallatae, Parapanteles aethiopicus, Parapanteles alternatus, Parapanteles aso, Parapanteles atellae, Parapanteles bagicha, Parapanteles cleo, Parapanteles cyclorhaphus, Parapanteles demades, Parapanteles endymion, Parapanteles epiplemicidus, Parapanteles expulsus, Parapanteles fallax, Parapanteles folia, Parapanteles furax, Parapanteles hemitheae, Parapanteles hyposidrae, Parapanteles indicus, Parapanteles javensis, Parapanteles jhaverii, Parapanteles maculipalpis, Parapanteles maynei, Parapanteles neocajani, Parapanteles neohyblaeae, Parapanteles nydia, Parapanteles prosper, Parapanteles prosymna, Parapanteles punctatissimus, Parapanteles regalis, Parapanteles sarpedon, Parapanteles sartamus, Parapanteles scultena, Parapanteles transvaalensis, Parapanteles turri, Parapanteles xanthopholis, Pholetesor acutus, Pholetesor brevivalvatus, Pholetesor extentus, Pholetesor ingenuoides, Pholetesor kuwayamai, Promicrogaster apidanus, Promicrogaster briareus, Promicrogaster conopiae, Promicrogaster emesa, Promicrogaster grandicula, Promicrogaster orsedice, Promicrogaster repleta, Promicrogaster typhon, Sathon bekilyensis, Sathon flavofacialis, Sathon laurae, Sathon mikeno, Sathon ruandanus, Sathon rufotestaceus, Venanides astydamia, Venanides demeter, Venanides parmula, and Venanides symmysta.
Metamorphosing insects often have complex and poorly known life histories. In particular, what they feed on during their larval stages remains unknown for the vast majority of species, and its documentation only results from difficult and time-intensive field observations, rearing or dissections. Through the application of a DNA analysis of gut contents in adult parasitoid wasps, we were able to selectively sequence a diagnostic DNA marker that permitted the identification of the host used by these wasps during their larval stages. By reproducing these results in species with different life histories, we excluded other potential sources of host DNA, confirming that after ingestion by the parasitoid larva the host DNA can persist through metamorphosis in the abdominal contents of the adult wasp. Our discovery considerably extends the applicability of molecular analysis of gut contents by enabling the documentation of food used by insects during their larval stages and thus increasing the accuracy and precision of food web studies. The 24% success rate of our approach is surprisingly high considering the challenging context for host DNA preservation, and we discuss the factors possibly affecting this rate. We propose molecular analysis of parasitoid linkages (MAPL) as a new method to document host-parasitoid associations at a faster pace and with unrivalled precision. Because of the key regulatory role of parasitoid wasps in ecosystems, which makes them the most commonly used biological control agents, MAPL will have immediate applications in both basic and applied biological sciences.
BackgroundComprehensive biotic surveys, or ‘all taxon biodiversity inventories’ (ATBI), have traditionally been limited in scale or scope due to the complications surrounding specimen sorting and species identification. To circumvent these issues, several ATBI projects have successfully integrated DNA barcoding into their identification procedures and witnessed acceleration in their surveys and subsequent increase in project scope and scale. The Biodiversity Institute of Ontario partnered with the rare Charitable Research Reserve and delegates of the 6th International Barcode of Life Conference to complete its own rapid, barcode-assisted ATBI of an established land trust in Cambridge, Ontario, Canada.New informationThe existing species inventory for the rare Charitable Research Reserve was rapidly expanded by integrating a DNA barcoding workflow with two surveying strategies – a comprehensive sampling scheme over four months, followed by a one-day bioblitz involving international taxonomic experts. The two surveys resulted in 25,287 and 3,502 specimens barcoded, respectively, as well as 127 human observations. This barcoded material, all vouchered at the Biodiversity Institute of Ontario collection, covers 14 phyla, 29 classes, 117 orders, and 531 families of animals, plants, fungi, and lichens. Overall, the ATBI documented 1,102 new species records for the nature reserve, expanding the existing long-term inventory by 49%. In addition, 2,793 distinct Barcode Index Numbers (BINs) were assigned to genus or higher level taxonomy, and represent additional species that will be added once their taxonomy is resolved. For the 3,502 specimens, the collection, sequence analysis, taxonomic assignment, data release and manuscript submission by 100+ co-authors all occurred in less than one week. This demonstrates the speed at which barcode-assisted inventories can be completed and the utility that barcoding provides in minimizing and guiding valuable taxonomic specialist time. The final product is more than a comprehensive biotic inventory – it is also a rich dataset of fine-scale occurrence and sequence data, all archived and cross-linked in the major biodiversity data repositories. This model of rapid generation and dissemination of essential biodiversity data could be followed to conduct regional assessments of biodiversity status and change, and potentially be employed for evaluating progress towards the Aichi Targets of the Strategic Plan for Biodiversity 2011–2020.
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