Major lineages of Nolidae (Lepidoptera, Noctuoidea) elucidated by molecular phylogenetics.

To elucidate the evolutionary relationships of the major lineages within the moth family Nolidae, we analysed a molecular dataset comprising eight independent gene regions (6.4 kbp), cytochrome c oxidase subunit I (COI) from the mitochondrial genome, and elongation factor-1α (EF-1α), ribosomal protein S5 (RpS5), carbamoylphosphate synthase domain protein (CAD), cytosolic malate dehydrogenase (MDH), glyceraldehyde-3-phosphate dehydrogenase (GAPDH), isocitrate dehydrogenase (IDH) and wingless genes from the nuclear genome, using parsimony and model-based evolutionary methods (maximum likelihood and Bayesian inference). Our analyses revealed a well-resolved phylogenetic hypothesis, again recovering the six previously recognized families within Noctuoidea (i.e. Oenosandridae, Notodontidae, Euteliidae, Erebidae, Nolidae and Noctuidae), and monophyly of the quadrifid Noctuoidea (i.e. Euteliidae, Erebidae, Nolidae and Noctuidae). The family Nolidae is diagnosed and characterized by two synapomorphies from morphology: construction of a ridged boat-shaped cocoon that bears a vertical exit slit at one end; and two other morphological character states: elongation of the forewing retinaculum into a bar-like or digitate condition and possession of a postpiracular counter-tympanal hood. We present a new phylogenetic hypothesis for Nolidae consisting of eight strongly supported subfamilies, two of which are erected here: Diphtherinae, Risobinae, Collomeninae subfam. nov., Beaninae subfam. nov., Eligminae, Westermanniinae, Nolinae and Chloephorinae. Where we are able, each monophyletic lineage is diagnosed by morphological autapomorphies and within each subfamily, monophyletic tribes and subtribes are circumscribed, most of which are also diagnosable by morphological apomorphies. We also describe two new taxa: Gelastocerini trib. nov. and Etannina subtrib. nov. The Neotropical subfamily Diphtherinae, here newly circumscribed, is considered to be the plesiomorphic sister lineage to the rest of Nolidae. Diphtherinae are characterized by loss of the proximal pair of metatibial spurs in males and by the presence of a frontal tubercle, which is presumably associated with a derived strategy of emergence from the cocoon.

Confirmed occurrence of the tribe Apameini (Lepidoptera, Noctuidae, Noctuinae) in the Neotropical region: a new genus endemic to Costa Rican montane forests.

The genus Nublapamea gen. nov. is described (type species: Tracheaaltivolans Schaus), here determined to belong to the primarily temperate Holarctic tribe Apameini (Noctuidae: Noctuinae). Currently known only from mid- to upper elevation montane forests of Costa Rica, Nublapamea is a disjunct southern extension of a largely northern hemisphere temperate region fauna. The life history of Nublapameaaltivolans is unknown; it may be associated with chusquea bamboo (Chusquea Kunth), as most Apameini are dietary specialists on graminoids.

A new Andean genus, Lafontaineana, with descriptions of four new species and two new Neotropical species of Panthea (Noctuidae, Pantheinae).

Lafontaineana Martinez, gen. nov. is proposed as a new Neotropical genus of Pantheinae, forming a sister group to Gaujonia Dognin, 1891 based on a phylogenetic analysis. In addition, one new combination and four new species are proposed: Lafontaineana marmorifera (Walker, 1865), comb. nov. (Colombia), Lafontaineana alexandrae Martinez, sp. nov. (Ecuador), Lafontaineana imama Martinez, sp. nov. (Colombia), Lafontaineana puma Martinez, sp. nov. (Ecuador), and Lafontaineana thuta Martinez, sp. nov. (Ecuador). Two new Neotropical species of Panthea are described, Panthea hondurensis Martinez, sp. nov. and Panthea taina Martinez, sp. nov.

Cryptic species among bumblebee mimics: an unrecognized Hemaris hawkmoth (Lepidoptera: Sphingidae) in eastern North America.

Through integrating molecular, morphological and natural history evidence, nominal Hemaris diffinis (Boisduval) of eastern North America is shown to include a second, cryptic species, Hemaris aethra (Strecker) stat. rev. Despite highly divergent mtDNA sequences and differing larval phenotypes, genitalic morphology, habitat and larval host plants, adults of H. aethra and sympatric H. diffinis are externally so similar that H. aethra has remained unrecognized for over a century. With a more northerly distribution than H. diffinis, H. aethra occurs from Manitoba to Nova Scotia and adjacent parts of the United States, the two species occurring in strict sympatry in eastern Ontario and likely other regions. Co-mimicry of Bombus Latreille bumblebee models has likely resulted in phenotypic convergence of H. diffinis and H. aethra, as the two do not appear to be sister taxa, the latter instead being more closely related to the western species H. thetis (Boisduval). The larvae of H. aethra are illustrated for the first time, together with diagnostic images and comparisons of adults. Lectotypes are designated for Hemaris tenuis Grote and Hemaris marginalis Grote.

Three species in one: a revision of Clemensiaalbata Packard (Erebidae, Arctiinae, Lithosiini).

Clemensiaalbata Packard, previously thought to be a single, widely distributed North American species, is here shown to consist of three distinct, primarily parapatric species: Clemensiaalbata Packard, C.umbrata Packard, stat. rev., and Clemensiaochreata Schmidt & Sullivan, sp. n. Adults and genitalic structures of the three species are illustrated.

A new species of Euchaetes Harris from southern Arizona (Erebidae, Arctiinae).

Euchaetesnancyae sp. n. is described from southeastern Arizona. Although superficially similar to species of Pygarctia Grote, structural and molecular variation shows it to be most closely related to Euchaeteshelena (Cassino). Adults, genitalic structure, eggs, and first instar larvae are described and illustrated. The larval host plant remains unknown. Euchaeteshelena is confirmed as occurring in Mexico.

A new species of Sympistis Hübner from Sapelo Island, Georgia, USA (Lepidoptera, Noctuidae, Oncocnemidinae).

A new species of the Sympistisbadistriga species-group, Sympistiseleaner Adams, sp. n. is described from Sapelo Island, a back-barrier island in coastal Georgia, United States of America. Adults and genitalia of S.eleaner are illustrated, in addition to adults of similar species in the Sympistisbadistriga species-group. The composition of this species-group is discussed.

A revision of Admetovis Grote, with the description of a new species from western North America (Noctuidae, Noctuinae, Hadenini).

The genus Admetovis Grote is revised. Admetovisicarus sp. n. is described from the mountains of western North America. A lectotype of Admetovisoxymorus Grote is designated. Illustrations of the adults, male and female genitalia, and distribution maps are presented, together with an identification key. The classification of the genus is reviewed resulting in its reassignment to the tribe Hadenini from Orthosiini.

Additions and corrections to the check list of the Noctuoidea (Insecta, Lepidoptera) of North America north of Mexico IV.

A summary of all taxonomic and nomenclatural changes to the check list of the Noctuoidea of North America north of Mexico since the last update published in 2015 is provided. A total of 64 changes are listed and discussed, consisting of 26 recently published changes and additions, and an additional 38 presented herein. One stat. n., one stat. rev., six syn. n., and two comb. n. are proposed for the first time. Orthimella Schmidt & Lafontaine nom. n. is proposed here as an objective replacement name for Himella Grote, 1874 [Noctuinae: Orthosiini], a junior homonym of Himella Dallas, 1852 [Hemiptera: Coreidae].

Probing planetary biodiversity with DNA barcodes: The Noctuoidea of North America.

This study reports the assembly of a DNA barcode reference library for species in the lepidopteran superfamily Noctuoidea from Canada and the USA. Based on the analysis of 69,378 specimens, the library provides coverage for 97.3% of the noctuoid fauna (3565 of 3664 species). In addition to verifying the strong performance of DNA barcodes in the discrimination of these species, the results indicate close congruence between the number of species analyzed (3565) and the number of sequence clusters (3816) recognized by the Barcode Index Number (BIN) system. Distributional patterns across 12 North American ecoregions are examined for the 3251 species that have GPS data while BIN analysis is used to quantify overlap between the noctuoid faunas of North America and other zoogeographic regions. This analysis reveals that 90% of North American noctuoids are endemic and that just 7.5% and 1.8% of BINs are shared with the Neotropics and with the Palearctic, respectively. One third (29) of the latter species are recent introductions and, as expected, they possess low intraspecific divergences.

What Azure blues occur in Canada? A re-assessment of Celastrina Tutt species (Lepidoptera, Lycaenidae).

The identity of Celastrina species in eastern Canada is reviewed based on larval host plants, phenology, adult phenotypes, mtDNA barcodes and re-assessment of published data. The status of the Cherry Gall Azure (Celastrina serotina Pavulaan & Wright) as a distinct species in Canada is not supported by any dataset, and is removed from the Canadian fauna. Previous records of this taxon are re-identified as Celastrina lucia (Kirby) and Celastrina neglecta (Edwards). Evidence is presented that both Celastrina lucia and Celastrina neglecta have a second, summer-flying generation in parts of Canada. The summer generation of Celastrina lucia has previously been misidentified as Celastrina neglecta, which differs in phenology, adult phenotype and larval hosts from summer Celastrina lucia. DNA barcodes are highly conserved among at least three North American Celastrina species, and provide no taxonomic information. Celastrina neglecta has a Canadian distribution restricted to southern Ontario, Manitoba, Saskatchewan and easternmost Alberta. The discovery of museum specimens of Celastrina ladon (Cramer) from southernmost Ontario represents a new species for the Canadian butterfly fauna, which is in need of conservation status assessment.

Century-Old DNA Barcodes Reveal Phylogenetic Placement of the Extinct Jamaican Sunset Moth, Urania sloanus Cramer (Lepidoptera: Uraniidae).

Analysis of the DNA barcode region of the cytochrome c oxidase 1 gene from a specimen of the extinct Jamaican sunset moth, Urania sloanus, places this species as a sister to the Central American U. fulgens. We found that all Urania F. species were closely related (<2.8% maximum divergence at COI), with the Cuban endemic U. boisduvalii appearing as sister to the rest. The low divergence in DNA barcodes and genitalic structures indicate that the Cuban U. poeyi and eastern Brazilian U. brasiliensis are geographic segregates of U. fulgens and U. leilus respectively, so the former two taxa are accordingly recognized as subspecies.

Revision of the Lacinipolia vicina (Grote) complex (Noctuidae, Noctuinae, Eriopygini).

The Lacinipolia vicina (Grote) species complex, previously consisting of Lacinipolia vicina, Lacinipolia teligera (Morrison), Lacinipolia pensilis (Grote), and Lacinipolia subalba Mustelin is revised to six species: Lacinipolia vicina (eastern USA), Lacinipolia teligera (southern Great Plains), Lacinipolia pensilis (Pacific Northwest and northern Rocky Mountains), Lacinipolia acutipennis (Grote), stat. rev. (= Lacinipolia subalba syn. n.) (western North America), Lacinipolia sareta (Smith), stat. rev. (Canada and western USA) and Lacinipolia dimocki, sp. n. (California and Pacific Northwest). Lectotypes are designated for Lacinipolia vicina, Lacinipolia teligera and Lacinipolia pensilis.

Additions and corrections to the check list of the Noctuoidea (Insecta, Lepidoptera) of North America north of Mexico III.

A total of 124 additions and corrections are listed and discussed for the check list of the Noctuoidea of North America north of Mexico published in 2010. Twenty-eight species are added to the list, 16 through new species descriptions, eight as a result of taxonomic splits, and four based on newly recorded species. Forty-eight species are deleted from the list, 41 through synonymy, and seven that were based on misidentifications. Twelve changes are corrections in the spelling of names, or changes in parentheses on dates of publication. Twenty-seven are changes in taxonomy of names where no species are added or deleted; eight changes involve the renumbering of existing species for better taxonomic arrangement. Within the text 2 stat. n., 10 stat. rev., 27 syn. n., 5 syn. rev., and 1 comb. n. are proposed for the first time.

Taxonomy and biogeography of the Nearctic Raphia Hübner (Lepidoptera, Noctuidae, Raphiinae).

The taxonomic status and biogeography of the North American Raphia species is reviewed using adult morphology, larval host plants, geographic phenotypic variation, and variation of mtDNA COI barcode sequences. Lack of diagnostic morphological differences, combined with relatively low mtDNA barcode divergences and clinal phenotypic variation in key geographic regions indicate that the six previously recognized species of North American Raphia are best interpreted as parapatric subspecies. Raphia frater abrupta Grote, stat. n., R. f. coloradensis Putnam-Cramer, stat. r., R. f. piazzi Hill, stat. n., and R. f. elbea Smith, stat. n., are accordingly revised to subspecies of R. frater Grote. Type locality restrictions are provided for Raphia abrupta and Raphia frater and a neotype is designated for Raphia frater var. coloradensis.

Polyphyly of Lichen-cryptic Dagger Moths: synonymy of Agriopodes Hampson and description of a new basal acronictine genus, Chloronycta, gen. n. (Lepidoptera, Noctuidae).

The taxonomic composition and systematic position of Agriopodes Hampson is examined through an integrated approach using adult and larval morphology, biology, and molecular sequence data. The type-species of Agriopodes, Moma fallax Herrich-Schäffer is shown to be derived within the Acronicta grisea Walker species-group; accordingly, Agriopodes is relegated to synonymy under Acronicta Ochsenheimer, syn. n. (Acronictinae). Additionally, molecular markers and morphology show that Agriopodes is not monophyletic: Agriopodes tybo (Barnes) is not closely related to A. fallax nor to Acronicta, and is transferred to a new genus, Chloronycta Schmidt & Anweiler, gen. n. The immature stages of Chloronycta tybo comb. n. are described and illustrated for the first time. Although previously treated as a valid species, we show that Agriopodes geminata (Smith) represents the northern terminus of clinal variation in wing pattern of A. fallax and synonymize A. geminata under A. fallax (syn. n.). The history and identity of Agriopodes corticosa (Boisduval), a nomen dubium, is discussed.

A transcontinental challenge--a test of DNA barcode performance for 1,541 species of Canadian Noctuoidea (Lepidoptera).

This study provides a first, comprehensive, diagnostic use of DNA barcodes for the Canadian fauna of noctuoids or "owlet" moths (Lepidoptera: Noctuoidea) based on vouchered records for 1,541 species (99.1% species coverage), and more than 30,000 sequences. When viewed from a Canada-wide perspective, DNA barcodes unambiguously discriminate 90% of the noctuoid species recognized through prior taxonomic study, and resolution reaches 95.6% when considered at a provincial scale. Barcode sharing is concentrated in certain lineages with 54% of the cases involving 1.8% of the genera. Deep intraspecific divergence exists in 7.7% of the species, but further studies are required to clarify whether these cases reflect an overlooked species complex or phylogeographic variation in a single species. Non-native species possess higher Nearest-Neighbour (NN) distances than native taxa, whereas generalist feeders have lower NN distances than those with more specialized feeding habits. We found high concordance between taxonomic names and sequence clusters delineated by the Barcode Index Number (BIN) system with 1,082 species (70%) assigned to a unique BIN. The cases of discordance involve both BIN mergers and BIN splits with 38 species falling into both categories, most likely reflecting bidirectional introgression. One fifth of the species are involved in a BIN merger reflecting the presence of 158 species sharing their barcode sequence with at least one other taxon, and 189 species with low, but diagnostic COI divergence. A very few cases (13) involved species whose members fell into both categories. Most of the remaining 140 species show a split into two or three BINs per species, while Virbia ferruginosa was divided into 16. The overall results confirm that DNA barcodes are effective for the identification of Canadian noctuoids. This study also affirms that BINs are a strong proxy for species, providing a pathway for a rapid, accurate estimation of animal diversity.

Comments on differences in classification of the superfamily Noctuoidea (Insecta, Lepidoptera) between Eurasia and North America.

An attempt is made to bring consensus between the classifications of the Noctuoidea in Europe and North America. Twelve points of disagreement between the check lists from the two regions are discussed and solutions recommended.

Lepidoptera family-group names proposed by Thaddeus William Harris in 1841.

In 1841, T. W. Harris published "A Report on the Insects of Massachusetts, Injurious to Vegetation," a seminal work in North American Entomology and one of the first New World publications to describe Lepidoptera species. Although appearing in several subsequent editions, the eight family-group names proposed by Harris were largely overlooked. In summarizing Harris' family-group names, we show that authorship for two Noctuidae names in current usage require changes: Acronictinae Harris, 1841 (originally as Acronyctadae) has priority over Heinemann, 1859, and Agrotini Harris, 1841 (as Agrotitidae) has priority over Rambur, 1848. Mamestradae Harris, 1841 is also a senior synonym of Mamestrinae Hampson, 1902 (Type genus: Mamestra Ochsenheimer, 1816), an available name that is currently a junior subjective synonym of Noctuidae: Noctuinae: Hadenini Guenée, 1837 (Type genus: Hadena Schrank, 1802). Geometridae: Ennominae: Hyberniini Harris, 1841 (as Hyberniadae), based on Erranis Hübner, [1825] (= Hybernia Berthold, 1827), has precedence over two family-group names with long-standing usage, Bistonini Stephens, 1850 and Boarmiini Duponchel, 1845, and a reversal of precedence for the latter two names over Hyberniini is proposed under ICZN guidelines.