DNA barcodes can be used to identify cryptic species of skipper butterflies previously detected by classic taxonomic methods and to provide first clues to the existence of yet other cryptic species. A striking case is the common geographically and ecologically widespread neotropical skipper butterfly Perichares philetes (Lepidoptera, Hesperiidae), described in 1775, which barcoding splits into a complex of four species in Area de Conservación Guanacaste (ACG) in northwestern Costa Rica. Three of the species are new, and all four are described. Caterpillars, pupae, and foodplants offer better distinguishing characters than do adults, whose differences are mostly average, subtle, and blurred by intraspecific variation. The caterpillars of two species are generalist grass-eaters; of the other two, specialist palm-eaters, each of which feeds on different genera. But all of these cryptic species are more specialized in their diet than was the morphospecies that held them. The four ACG taxa discovered to date belong to a panneotropical complex of at least eight species. This complex likely includes still more species, whose exposure may require barcoding. Barcoding ACG hesperiid morphospecies has increased their number by nearly 10%, an unexpectedly high figure for such relatively well known insects.
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 Conservacion de Guanacaste (ACG), northwestern Costa Rica. This provides the world's best location-based dataset for studying the taxonomy and host relationships of caterpillar parasitoids. Among Hymenoptera, Microgastrinae (Braconidae) is the most diverse and commonly encountered parasitoid subfamily, with many hundreds of species delineated to date, almost all undescribed. Here, we reassess the limits of the genus Apanteles sensu stricto, describe 186 new species from 3,200+ parasitized caterpillars of hundreds of ACG Lepidoptera species, and provide keys to all 205 described Apanteles from Mesoamerica -including 19 previously described species in addition to the new species. The Mesoamerican Apanteles are assigned to 32 species-groups, all but two of which are newly defined. Taxonomic keys are presented in two formats: traditional dichotomous print versions and links to electronic interactive versions (software Lucid 3.5). Numerous illustrations, computer-generated descriptions, distributional information, wasp biology, and DNA barcodes (where available) are presented for every species. All morphological terms are detailed and linked to the Hymenoptera Anatomy Ontology website. DNA barcodes (a standard fragment of the cytochrome c oxidase I (COI) mitochondrial gene), information on wasp biology (host records, solitary/gregariousness of wasp larvae), ratios of morphological features, and wasp microecological distributions were used to help clarify boundaries between morphologically cryptic species within species-complexes. Because of the high accuracy of host identification for about 80% of the wasp species studied, it was possible to analyze host relationships at a regional level. The ACG species of Apanteles attack mainly species of Hesperiidae, Elachistidae and Crambidae (Lepidoptera). About 90% of the wasp species with known host records seem to be monophagous or oligophagous at some level, parasitizing just one host family and commonly, just one species of caterpillar. Only 15 species (9%) parasitize species in more than one family, and some of these cases are likely to be found to be species complexes. We have used several information sources and techniques (traditional taxonomy, molecular, software-based, biology, and geography) to accelerate the process of finding and describing these new species in a hyperdiverse group such as Apanteles. The following new taxonomic and nomenclatural acts are proposed. Four species previously considered to be Apanteles are transferred to other microgastrine genera: Dolichogenidea hedyleptae (Muesebeck, 1958), comb. n., Dolichogenidea politiventris (Muesebeck, 1958), comb. n., Rhygoplitis sanctivincenti (Ashmead, 1900), comb. n., and Illidops scutellaris (Muesebeck, 1921), comb. rev. One European species that is a secondary homonym to a Mesoamerican species is removed from Apanteles and transferred to another genus: Iconella albinervis (Tobias, 1964), stat. rev. The name Apanteles albinervican Shenefelt, 1972, is an invalid replacement name for Apanteles albinervis (Cameron, 1904), stat. rev., and thus the later name is reinstated as valid. The following 186 species, all in Apanteles and all authored by Fernandez-Triana, are described as species nova: adelinamoralesae, adrianachavarriae, adrianaguilarae, adrianguadamuzi, aichagirardae, aidalopezae, albanjimenezi, alejandromasisi, alejandromorai, minorcarmonai, alvarougaldei, federicomatarritai, anabellecordobae, rostermoragai, anamarencoae, anamartinesae, anapiedrae, anariasae, andreacalvoae, angelsolisi, arielopezi, bernardoespinozai, bernyapui, bettymarchenae, bienvenidachavarriae, calixtomoragai, carloscastilloi, carlosguadamuzi, eliethcantillanoae, carlosrodriguezi, carlosviquezi, carloszunigai, carolinacanoae, christianzunigai, cinthiabarrantesae, ciriloumanai, cristianalemani, cynthiacorderoae, deifiliadavilae, dickyui, didiguadamuzi, diegoalpizari, diegotorresi, diniamartinezae, duniagarciae, duvalierbricenoi, edgarjimenezi, edithlopezae, eduardoramirezi, edwinapui, eldarayae, erickduartei, esthercentenoae, eugeniaphilipsae, eulogiosequeira, felipechavarriai, felixcarmonai, fernandochavarriai, flormoralesae, franciscopizarroi, franciscoramirezi, freddyquesadai, freddysalazari, gabrielagutierrezae, garygibsoni, gerardobandoi, gerardosandovali, gladysrojasae, glenriverai, gloriasihezarae, guadaluperodriguezae, guillermopereirai, juanmatai, harryramirezi, hectorsolisi, humbertolopezi, inesolisae, irenecarrilloae, isaacbermudezi, isidrochaconi, isidrovillegasi, ivonnetranae, jairomoyai, javiercontrerasi, javierobandoi, javiersihezari, jesusbrenesi, jesusugaldei, jimmychevezi, johanvargasi, jorgecortesi, jorgehernandezi, josecalvoi, josecortesi, josediazi, josejaramilloi, josemonteroi, joseperezi, joserasi, juanapui, juancarrilloi, juangazoi, juanhernandezi, juanlopezi, juanvictori, juliodiazi, juniorlopezi, keineraragoni, laurahuberae, laurenmoralesae, leninguadamuzi, leonelgarayi, lilliammenae, lisabearssae, luciariosae, luisbrizuelai, luiscanalesi, luiscantillanoi, luisgarciai, luisgaritai, luishernandezi, luislopezi, luisvargasi, manuelarayai, manuelpereirai, manuelriosi, manuelzumbadoi, marcobustosi, marcogonzalezi, marcovenicioi, mariachavarriae mariaguevarae, marialuisariasae, mariamendezae, marianopereirai, mariatorrentesae, sigifredomarini, marisolarroyoae, marisolnavarroae, marvinmendozai, mauriciogurdiani, milenagutierrezae, monicachavarriae, oscarchavesi, osvaldoespinozai, pablotranai, pabloumanai, pablovasquezi, paulaixcamparijae, luzmariaromeroae, petronariosae, randallgarciai, randallmartinezi, raulacevedoi, raulsolorsanoi, wadyobandoi, ricardocaleroi, robertmontanoi, robertoespinozai, robertovargasi, rodrigogamezi, rogerblancoi, rolandoramosi, rolandovegai, ronaldcastroi, ronaldgutierrezi, ronaldmurilloi, ronaldnavarroi, ronaldquirosi, ronaldzunigai, rosibelelizondoae, ruthfrancoae, sergiocascantei, sergioriosi, tiboshartae, vannesabrenesae, minornavarroi, victorbarrantesi, waldymedinai, wilbertharayai, williamcamposi, yeissonchavesi, yilbertalvaradoi, yolandarojasae, hazelcambroneroae, zeneidabolanosae.
Background: Skipper butterflies (Hesperiidae) are a relatively well-studied family of Lepidoptera. However, a combination of DNA barcodes, morphology, and natural history data has revealed several cryptic species complexes within them. Here, we investigate three DNA barcode lineages of what has been identified as Urbanus belli (Hesperiidae, Eudaminae) in Area de Conservacion Guanacaste (ACG), northwestern Costa Rica. Results: Although no morphological traits appear to distinguish among the three, congruent nuclear and mitochondrial lineage patterns show that "Urbanus belli" in ACG is a complex of three sympatric species. A single strain of Wolbachia present in two of the three cryptic species indicates that Urbanus segnestami Burns (formerly Urbanus belliDHJ01), Urbanus bernikerni Burns (formerly Urbanus belliDHJ02), and Urbanus ehakernae Burns (formerly Urbanus belliDHJ03) may be biologically separated by Wolbachia, as well as by their genetics. Use of parallel sequencing through 454-pyrosequencing improved the utility of ITS2 as a phylogenetic marker and permitted examination of the intra-and interlineage relationships of ITS2 variants within the species complex. Interlineage, intralineage and intragenomic compensatory base pair changes were discovered in the secondary structure of ITS2. Conclusion: These findings corroborate the existence of three cryptic species. Our confirmation of a novel cryptic species complex, initially suggested by DNA barcode lineages, argues for using a multi-marker approach coupled with next-generation sequencing for exploration of other suspected species complexes.
90 species of Euplectrus are treated: 55 newly described, all from Area de Conservación Guanacaste (ACG), and 35 previously described species, of which 20 occur in ACG. Three of the previously described species (Euplectrusbrasiliensis Ashmead, Euplectrushircinus (Say), Euplectrusronnai (Brèthes)) have unknown status, owing to missing or severely damaged type material. The new species, all authored by C. Hansson, are: Euplectrusalejandrovalerioi, Euplectrusalexsmithi, Euplectrusalvarowillei, Euplectrusandybennetti, Euplectrusandydeansi, Euplectrusannettewalkerae, Euplectrusbillbrowni, Euplectrusbobwhartoni, Euplectruscarlosarmientoi, Euplectruscarlrettenmeyeri, Euplectruscharlesmicheneri, Euplectruscharlesporteri, Euplectruschrisdarlingi, Euplectruschrisgrinteri, Euplectruscorriemoreauae, Euplectrusdaveroubiki, Euplectrusdavesmithi, Euplectrusdavidwahli, Euplectrusdianariasae, Euplectrusdonquickei, Euplectruseowilsoni, Euplectrusgarygibsoni, Euplectrusgavinbroadi, Euplectrusgerarddelvarei, Euplectrushenrytownesi, Euplectrushowelldalyi, Euplectrushugokonsi, Euplectrusiangauldi, Euplectrusjacklonginoi, Euplectrusjesusugaldei, Euplectrusjimwhitfieldi, Euplectrusjjrodriguezae, Euplectrusjohnheratyi, Euplectrusjohnlasallei, Euplectrusjohnnoyesi, Euplectrusjosefernandezi, Euplectruslubomirmasneri, Euplectrusmarkshawi, Euplectrusmikegatesi, Euplectrusmikeschauffi, Euplectrusmikesharkeyi, Euplectrusninazitaniae, Euplectruspammitchellae, Euplectruspaulhansoni, Euplectruspaulheberti, Euplectruspaulhurdi, Euplectrusphilwardi, Euplectrusrobbinthorpi, Euplectrusronaldzunigai, Euplectrusroysnellingi, Euplectrusscottshawi, Euplectrussondrawardae, Euplectrussydneycameronae, Euplectrusvictoriapookae, Euplectruswonyoungchoi. The species are described or redescribed, and thoroughly and uniformly illustrated, and included in two identification keys, one for females and one for males. Lectotypes are designated for eight species: Euplectruscatocalae Howard (♂), Euplectrusjunctus Gahan (♀), Euplectrusleucotrophis Howard (♂), Euplectrusmarginatus Ashmead (♀), Euplectruspachyscaphus Girault (♀), Euplectrusplatyhypenae Howard (♂), Euplectrussemimarginatus Girault (♀), Heteroscapusronnai Brèthes (♂). One synonym is established: Euplectruswalteri Schauff is a junior synonym of Euplectrustestaceipes (Cameron). Brief image notes and host records are provided on the natural history of the wasps as well as the details of their morphology. Hosts are known for 74 Euplectrus species.
Five species of Trigonalidae, hyperparasitoids of Ichneumonidae (Hymenoptera) and Tachinidae (Diptera) that parasitize caterpillars (Lepidoptera), have been reared during the ongoing caterpillar inventory of Area de Conservacion Guanacaste (ACG), Guanacaste Province, northwestern Costa Rica: Lycogaster apicipennis (Cameron), Taeniogonalos woodorum Smith, sp. n., Taeniogonalos fasciatipennis (Cameron), Trigonalys championi Cameron, and Trigonalys maculifrons Sharp. Morphological and DNA barcoding data support species separation of these generalist hyperparasitoids. Taeniogonalos gundlachii (Cresson) is not a widespread, color-variable species as previously treated and is probably confined to eastern North America. The species previously considered as T gundlachii in Costa Rica is regarded as Taeniogonalos fasciatipennis, a species found only in ACG dry forest. Taeniogonalos woodorum is a similar species but found only in the ACG rain forest. Habitat and host records are given for these five species of trigonalids.
We discuss 45 Costa Rican species of Ethmia Hubner, 1819 including 23 previously described: E. delliella (Fernald), E. bittenella (Busck), E. festiva Busck, E. scythropa Walsingham, E. perpulchra Walsingham, E. terpnota Walsingham, E. elutella Busck, E. janzeni Powell, E. ungulatella Busck, E. exornata (Zeller), E. phylacis Walsingham, E. mnesicosma Meyrick, E. chemsaki Powell, E. baliostola Walsingham, E. duckworthi Powell, E. sandra Powell, E. nigritaenia Powell, E. catapeltica Meyrick, E. lichyi Powell, E. transversella Busck, E. similatella Busck, E. hammella Busck, E. linda Busck, and 22 new species: E. blaineorum, E. millerorum, E. dianemillerae, E. adrianforsythi, E. stephenrumseyi, E. berndkerni, E. dimauraorum, E. billalleni, E. ehakernae, E. helenmillerae, E. johnpringlei, E. laphamorum, E. petersterlingi, E. lesliesaulae, E. turnerorum, E. normgershenzi, E. nicholsonorum, E. hendersonorum, E. randyjonesi, E. randycurtisi, E. miriamschulmanae and E. tilneyorum. We illustrate all species and their male and female genitalia, along with distribution maps of Costa Rican localities. Immature stages are illustrated for 11 species, and food plants are listed when known. Gesneriaceae is added as a new food plant family record for Ethmia. CO1 nucleotide sequences ("DNA barcodes") were obtained for 41 of the species.
La Región del Biobío en el centro sur de Chile es una zona de transición climática donde conviven bosques templados y esclerófilos, generando una zona de alta diversidad. Sin embargo, el área ha sido fuertemente afectada por la intervención antrópica y solo quedan algunos relictos de matorral y bosque nativo. Los geométridos, al igual que muchos insectos, están estrechamente asociados a la vegetación y por lo tanto serán afectados directamente por la intervención que se efectúe sobre esta. Así, en este estudio evaluamos los patrones de distribución y diversidad de geométridos en la Región del Biobío, con el objetivo de proponer sitios de alta prioridad para conservación mediante la identificación de áreas de endemismo y hot spots de diversidad. Los datos disponibles fueron procesados mediante análisis de parsimonia de endemismo y análisis de complementariedad, apoyados por SIG para rellenar vacíos y extrapolar la distribución de algunas especies de acuerdo a criterios establecidos. Encontramos una diversidad de 120 especies de geométridos, que corresponden a un 37.5 % de la diversidad nacional, distribuidas en seis áreas de endemismo, localizadas en las formaciones vegetacionales más frecuentes en la Región del Biobío: bosque alto-montano de Nahuelbuta, bosque esclerófilo de los arenales, bosque caducifolio de Concepción, bosque caducifolio andino del Biobío y bosque caducifolio de la frontera. El análisis de complementariedad determinó que 18 celdas contienen el número total de especies en la región. Basado en los análisis anteriores, se determinó que hay cinco áreas que deben ser consideradas como sitios de alta prioridad para la conservación de geométridos: (1) la ladera occidental de la Cordillera de Nahuelbuta y sector costero adyacente, (2) el área Pencopolitana de la zona litoral de la región, (3) Cerro Negro-Quillón, (4) Las Trancas, y (5) la zona del valle del Queuco-Alto Biobío.The Biobio Region in south-central Chile is a climatic transition area where temperate and sclerophyllous forests cooccur, generating a high diversity zone. However, the area has been strongly affected by antropic intervention and only a few relicts of native forest and shrubland are left. Geometrid moths, like many other insects, are closely associated to the vegetation and therefore they will be directly affected by antropic intervention. Thus, in this study we assessed the patterns of distribution and diversity of geometrids in the Biobio Region, aiming to propose high-priority sites for conservation by identifying endemism areas and diversity hot spots. The available data were processed by parsimony analysis of endemism and complementarity analysis, helped by GIS, for to fill up records and to extrapolate the distribution of some species according to established criteria. We found a diversity of 120 geometrid species, corresponding to 37.5 % of the Chilean diversity, distributed in six endemism areas located in the most frequent vegetational formations in the Biobio region: the Nahuelbuta high-mountain forest, the sclerophyllous forest of sandy grounds, the deciduous forest of Concepción, the Andean deciduous forest of the Biobío, and the deciduous forest of the frontier. Complementarity analysis revealed that there were 18 square plots which included the total number of species in the region. Based on the above analyses, we determined that five areas should be considered as high-priority sites for the conservation of geometrids: (1) the western slope of the Nahuelbuta mountain range and its adjacent coastal sector, (2) the Pencopolitan area of the coastal zone of the region, (3) Cerro Negro-Quillón, (4) Las Trancas, and (5) the Alto Biobío-Queuco valley zone.
Twelve species of Costa Rican Lytopylus are treated; these include all species reared from Lepidoptera caterpillars in Area de Conservación Guanacaste, Costa Rica, over 32 years of caterpillar inventory, as well as two species recorded in the literature as occurring in Costa Rica. Ten new species are described, i.e., Lytopylus bradzlotnicki, Lytopylus colleenhitchcockae, Lytopylus gregburtoni, Lytopylus jessicadimauroae, Lytopylus jessiehillae, Lytopylus mingfangi, Lytopylus rebeccashapleyae, Lytopylus robpringlei, Lytopylus sandraberriosae, Lytopylus vaughntani. The following species are transferred to Lytopylus: Metriosoma flavicalcar Enderlein 1920 to Lytopylus flavicalcarcomb. n.; Bassus macadamiae Briceño and Sharkey 2000 to Lytopylus macadamiaecomb. n.; Metriosoma bicarinatum Enderlein 1920 to Lytopylus bicarinatumcomb. n.; Metriosoma brasiliense Enderlein 1920 to Lytopylus brasiliensecomb. n.; Bassus tayrona Campos 2007 to Lytopylus tayronacomb. n.; Microdus femoratus Cameron 1887 to Lytopylus femoratuscomb. n.; Microdus melanocephalus Cameron 1887 to Lytopylus melanocephaluscomb. n.; Bassus pastranai Blanchard 1952 to Lytopylus pastranaicomb. n.; Agathis nigrobalteata Cameron 1911 to Lytopylus nigrobalteatuscomb. n. Two keys to species of Lytopylus are presented, one interactive and the other static.
Although central to much biological research, the identification of species is often difficult. The use of DNA barcodes, short DNA sequences from a standardized region of the genome, has recently been proposed as a tool to facilitate species identification and discovery. However, the effectiveness of DNA barcoding for identifying specimens in species-rich tropical biotas is unknown. Here we show that cytochrome c oxidase I DNA barcodes effectively discriminate among species in three Lepidoptera families from Area de Conservación Guanacaste in northwestern Costa Rica. We found that 97.9% of the 521 species recognized by prior taxonomic work possess distinctive cytochrome c oxidase I barcodes and that the few instances of interspecific sequence overlap involve very similar species. We also found two or more barcode clusters within each of 13 supposedly single species. Covariation between these clusters and morphological and/or ecological traits indicates overlooked species complexes. If these results are general, DNA barcoding will significantly aid species identification and discovery in tropical settings.