Lasionycta Aurivillius 1892
Creators: name: Crabo, Lars
name: Lafontaine, Donald
Publisher: name: Zenodo
Year: 2009
Subjects: Biodiversity
Taxonomy
Animalia
Arthropoda
Insecta
Lepidoptera
Noctuidae
Lasionycta

description: <i>Lasionycta</i> Aurivillius <i>Lasionycta</i> Aurivillius, 1892b: 285. Type species: <i>Phlogophora skraelingia</i> Herrich- Schäffer by original designation. <i>Lasiestra</i> Hampson, 1905: 47. Type species: <i>Dianthoecia phoca</i> Möschler by original designation. <i>Pseudanarta</i> Kozhanchikov, 1947: 18. Type species: <i>Anarta bohemani</i> Staudinger by original designation. Preoccupied by <i>Pseudanarta</i> Grote, 1878. <i>Anartomima</i> Boursin, 1952: 55. Replacement name for <i>Pseudanarta</i> Kozhanchikov. <b>Diagnosis.</b> <i>Lasionycta</i> is placed in Hadeninae based on trifine hindwing venation (vein M2 vestigial), hairy eyes, and larva with a dorsoventrally flattened spinneret with the opening shaped as a narrow transverse slit with an apical fringe. Within Hadeninae, <i>Lasionycta</i> is classified in Eriopygini Fibiger and Lafontaine based on an undivided larval hypopharynx, lack of an inner tooth on the larval mandible, and features of the larval spinneret, including elongated shape and apical fringe confined to the end of groove (Fibiger and Lafontaine 2005). Within Eriopygini, <i>Lasionycta</i> is characterized by the male genitalia, which have the following features: 1) an uncus with a nearly cylindrical base with a pointed or dorso-ventrally flattened tip; 2) a strap-like valve bearing a C-shaped clasper, an elongate digitus oriented 30–90° to the axis, a small to moderately large cucullus with a simple or compound corona, and a membranous flap on the mesial surface distal to the sacculus (post-saccular flap); 3) an elongate coiled or spiraled vesica bearing zero to six pointed subbasal cornuti with broad bases and a distal field of minute cornuti or one or two rows of spine-like cornuti oriented perpendicular to the vesica axis. The distal cornuti field is adjacent to a membranous wrinkled band running the length of the vesica. <b>Description. Head <i>–</i></b> Male antenna beadlike, biserrate, or bipectinate (1.1 <i>–</i> 5.4× as wide as central shaft) and fasciculate. Female antenna filiform and ciliate. Scape with a tuft of longer scales at dorsal base of antenna. Eye hairy, round to ellipsoid. Head covered in hair-like scales. <b>Thorax <i>–</i></b> Vestiture dense hair-like scales or mixture of hair-like and weakly spatulate scales. Foretibia with epiphysis. Tarsal segments with three rows of spine-like setae. <b>Wings <i>–</i></b> Ground color light to dark gray or gray brown. Wings of most species with typical noctuine lines and spots present (indistinct in several species). Hindwing nearly uniform gray or lighter gray, white, or yellow with variable dark suffusion and contrasting darker discal spot, postmedial line, and marginal band. <b>Male genitalia <i>–</i></b> Uncus curved, slightly laterally compressed with hook-like apex or with distal portion dorsoventrally flattened with ovoid or spatulate apex. Valve elongate, strap-like, 4.5–8.0× as long as wide. Sacculus rounded, relatively weak, with costal lobe extending below or above costal margin of valve. Valve bearing a membranous flap on ventromesial surface distal to sacculus ("postsaccular flap") (lacking in <i>L</i>. <i>secedens</i> species-group). Distal portion of valve weakly to moderately constricted below cucullus (neck). Cucullus 0.65–1.50× as wide as valve, with corona of 1–4 rows of stout claw-like setae. Clasper cylindrical and curved posterodorsad. Digitus moderately sclerotized, oriented 30–45° ventrad, finger-like with variable width base, or more heavily sclerotized, oriented 90° ventrad, with a basal flange or prong (<i>L</i>. <i>subdita</i> species-group). Aedeagus cylindrical, tubular, or with single thorn-like or multiple-tipped raised apical projection on right; vesica 1.5–3.0× as long as aedeagus, curved and twisted to nearly straight beyond a subbasal bend or coil, distal part extending right and ventral to distal aedeagus, with zero to six basal cornuti (variable between and within species) comprised of a central spike from a smooth to crenulate button-like base; vesica with a wrinkled lightly sclerotized band along entire length, distal vesica with adjacent field of multiple setae as a band of in- numerable velvety setae or one or more rows of stouter setae oriented perpendicular to vesica axis. <b>Female genitalia <i>–</i></b> Ovipositor lobe membranous and pad-like covered with hair-like setae or stout and conical covered with short stout setae. Abdominal segment VIII 0.3–1.2× as long as wide, short in species with pad-like ovipositors and elongate in species with conical ovipositors, covered with variable number of short setae, greatest ventrally and along posterior margin. Ductus bursae moderately sclerotized, cylindrical to dorsoventrally flattened, distal ventral aspect turned onto adjacent dorsal membranous bursa as a flange-like supporting structure; some species groups with posterior end heavily sclerotized and expanded to left (<i>L</i>. <i>secedens</i> species-group) or anterior end expanded to right (<i>L</i>. <i>subdita</i> species-group). Corpus bursae membranous, spherical or ovoid, with 3 elongate signa separated by 90°, one on each surface except side with appendix bursae (1 ventral signum in <i>L</i>. <i>secedens</i> species-group), with very weak to prominent mesial constriction, most pronounced on dorsal side near appendix bursae origin. Appendix bursae membranous or lightly sclerotized, arising from left side (<i>L</i>. <i>skraelingia</i> species-group) or dorsum of posterior corpus bursae and extending anteriorly to left, or posterodorsad from corpus bursae; shape of appendix weakly to strongly coiled 570° (<i>L</i>. <i>phaea</i> and <i>L</i>. <i>secedens</i> species-groups, respectively) or straight with subapical 90° bend. Ductus seminalis at distal appendix bursae (posterior end in most species, anterior end in <i>L</i>. <i>skraelingia</i> species-group). <b>Early stages.</b> Larvae are known for <i>L</i>. <i>skraelingia</i> (Herrich-Schäffer), <i>L</i>. <i>secedens</i> (Walker), <i>L</i>. <i>leucocycla</i> (Staudinger), <i>L</i>. <i>staudingeri</i> (Aurivillius), and <i>L</i>. <i>perplexa</i> (Smith) (Godfrey 1972; Ahola and Silvonen 2008). Hypopharynx undivided, mandible lacking a tooth on inner surface, spinneret long and narrow, and fringe at spinneret apex confined to a small area at end of dorsal groove (Fibiger and Lafontaine 2005). <b>Remarks.</b> The <i>Lasionycta</i> species in this revision are included in <i>Anartomima</i> Boursin, <i>Lasiestra</i> Hampson, and <i>Lasionycta</i> in the Moths of North America checklist (Franclemont and Todd 1983). <i>Lasiestra</i> (type species: <i>Dianthoecia phoca</i> Möschler) has since been synonymized with <i>Lasionycta</i> (Lafontaine et al. 1986). <i>Lasionycta secedens</i> was first treated as a <i>Lasionycta</i> in Europe (Hacker et al. 2002). It resembles <i>L</i>. <i>skraelingia</i> and <i>L</i>. <i>phaea</i> (Hampson) and like them the vesica is long; superficially it differs mainly in hindwing color. Its generic placement was scrutinized for this work because both sexes of <i>L</i>. <i>secedens</i> differ from other <i>Lasionycta</i> by several seemingly unique anatomic features. Characters intermediate between <i>L</i>. <i>secedens</i> and other <i>Lasionycta</i> species were found in the <i>L</i>. <i>phaea</i> and <i>L</i>. <i>mutilata</i> species-groups. Also, CO1 gene sequence distance analysis nests <i>L</i>. <i>secedens</i> among other <i>Lasionycta</i> speciesgroups, usually closest to the <i>L</i>. <i>phaea</i> species-group. This supports its inclusion in <i>Lasionycta</i>, and it is herein treated as the sole member of the <i>L</i>. <i>secedens</i> species-group. A check list of the North American Lasionycta species is presented in Table 2. Species included in <i>Lasionycta</i> in recent European literature are more diverse in appearance, morphology, and biology than in North America. Hacker et al. (2002) list 18 Palaearctic species in the genus. Four of these (<i>L</i>. <i>skraelingia</i>, <i>L</i>. <i>leucocycla</i>, <i>L</i>. <i>staudingeri</i>, and <i>L</i>. <i>secedens</i>) are Holarctic and are included in the current revision. Two other Asian species, <i>L</i>. <i>buraetica</i> Kononenko and <i>L</i>. <i>corax</i> Kononenko, are related to Nearctic <i>L</i>. <i>phaea</i>, as is <i>L</i>. <i>alpicola</i> Lafontaine &amp; Kononenko (Lafontaine and Kononenko 1988) which was omitted from this list. Four other species, <i>proxima</i>, <i>calberlai</i> (Staudinger), <i>imbecilla</i> (Fabricius), and <i>impar</i> (Staudinger), occur in Europe and were described and illustrated by Hacker et al. (2002). Th ey have no close relatives in North America and were therefore evaluated to determine their relationship to other <i>Lasionycta</i>. <i>Proxima</i>, the type species of <i>Lasionhada</i> Berio, 1981, has genitalia (Figs 190 and 245) similar to other <i>Lasionycta</i>. Th e male genital capsule resembles closely those of the <i>Lasionycta leucocycla</i> species-group in all characters including the presence of the post-saccular flap. However, the vesica differs from those of <i>Lasionycta</i> species in this revision in that the elongate band of distal setae are positioned on the membranous band, not at its margin. Th e female genitalia are similar to those of the <i>L</i>. <i>leucocycla</i> species-group but lack signa on the corpus bursae and have long symmetrically pointed ovipositor lobes, characters not found elsewhere in <i>Lasionycta</i>. Th e CO1 sequence of <i>proxima</i> suggests that it is an outlier from other <i>Lasionycta</i> species, usually clustering adjacent to them but occasionally grouping with related genera like <i>Tricholita</i> Grote (Fig. 247). It also differs biologically from other <i>Lasionycta</i> in that it occurs in dry temperate areas as low as sea level, whereas other <i>Lasionycta</i> species prefer conifer forests or tundra. For these reasons we transfer <i>proxima</i> back to <i>Lasionhada</i> as <i>Lasionhada proxima</i> (Hübner), comb. rev. Th e morphologic and DNA data suggest that <i>Lasionhada</i> is the sister group to <i>Lasionycta</i>. <i>Calberlai</i> is the type species of <i>Clemathada</i> Beck, 1991 and was moved from <i>Lasionycta</i> back to this genus by Fibiger and Lafontaine (2005). Th ey noted that it belongs in Hadenini, not Eriopygini, based on larva characters including a divided hypopharynx. <i>Imbecilla</i> is the type species of <i>Eriopygodes</i> Hampson, 1905 and was first associated with <i>Lasionycta</i> by Hacker et al. (2002). It was examined and differs significantly from <i>Lasionycta</i> in several respects, including presence of a stout broadly ovate clasper, a vestigial digitus, and no postsaccular flap of the male valve (Fig. 191). It does not group with other <i>Lasionycta</i> on CO1 sequence analysis, instead clustering with <i>Lacinipolia lorea</i> (Guenée), which it also resembles superficially. <i>Lacinipolia</i> McDunnough remains a large heterogeneous genus in need of revision. <i>Imbecilla</i> is herein transferred back to <i>Eriopygodes</i> as <i>E</i>. <i>imbecilla</i> (Fabricius), comb. rev. <i>Impar</i> was not available for examination for this revision but is illustrated by Hack- er et al. (2002). Its habitus and vesica are somewhat atypical for <i>Lasionycta</i> although the genitalia are similar. We retain it in <i>Lasionycta</i> pending further review of the Eurasian Eriopygini. The other eight species in the Hacker list (<i>L</i>. <i>hospita</i> Bang-Haas, <i>L</i>. <i>melanographa</i> Varga, <i>L</i>. <i>montanoides</i> Poole, <i>L</i>. <i>orientalis</i> (Alpharéky), <i>L</i>. <i>decreta</i> (Püngeler), <i>L</i>. <i>draudti</i> (F. Wagner), <i>L</i>. <i>hampsoni</i> Varga, and <i>L</i>. <i>levicula</i> (Püngeler)) are Asian and were not otherwise evaluated by Hacker et al. (2002) or by us in the current work. Only the first four of these species are included in <i>Lasionycta</i> by Poole (1989). Four North American species included in <i>Lasionycta</i> in recent checklists (Franclemont and Todd 1982; Poole 1989) (<i>L</i>. <i>insolens</i> (Grote), <i>L</i>. <i>arietis</i> (Grote), <i>L</i>. <i>wy- atti</i> (Barnes &amp; Benjamin), and <i>L</i>. <i>ochracea</i> (Smith) and the recently described <i>L</i>. <i>sala</i> Troubridge &amp; Mustelin) occur on Pacific Ocean sand beaches. Th eir larvae and adults differ structurally from other <i>Lasionycta</i>. Godfrey (1972) grouped <i>L</i>. <i>arietis</i> and <i>L</i>. <i>wyatti</i> separately from other <i>Lasionycta</i> (<i>L</i>. <i>perplexa</i> and an identified species of “ <i>Lasiestra</i> ” from Mt Rainier) in a comprehensive review of North American Hadenine larvae, recognizing that they formed a “very unique group.” These beach species cluster on CO1 distance analysis, consistently nesting among <i>Lacinipolia</i> species and removed from other <i>Lasionycta</i>. Th ese data indicate that they are not closely related to <i>Lasionycta</i>. A new genus, <i>Psammopolia</i> Crabo &amp; Lafontaine, is described for them in this paper. Many <i>Lasionycta</i> are alpine and have virtually no prospect for gene exchange between populations in different mountain ranges. It is not surprising that many populations differ slightly in appearance from those from nearby ranges. This revision required many decisions on whether to treat various levels of dissimilar disjunct populations as species, subspecies, or variation. After discovering convincing evidence that several <i>L</i>. <i>leucocycla</i> species-group taxa previously considered subspecies of <i>L</i>. <i>leucocycla</i> were species (e.g., <i>L</i>. <i>poca</i> (Barnes &amp; Benjamin) and <i>L</i>. <i>flanda</i> (Smith)) greater weight was given to superficial characters of pattern and color, taking care not to over interpret variability in species like <i>L</i>. <i>uniformis</i> (Smith). Proposed changes were tested against DNA data to formulate the most congruous final classification. Nonetheless, we usually gave greater weight to morphology and habitus than DNA and recognize several species without any DNA sequence differences, especially where the ranges were found to abut or overlap without any intermediate forms in the areas of sympatry. Taxa were treated as subspecies when there were significant differences in appearance between large regional populations with only slight overlap in characters, especially in species with multiple CO1 haplotypes in which barcode differences were considered to be less reliable than in groups with less variable DNA. We resisted the temptation to name small populations (e.g., <i>L</i>. <i>promulsa</i> (Morrison) from the Uinta and Wasatch Mountains), except when highly disjunct (<i>L</i>. <i>uniformis shasta</i> ssp. n. in California and <i>L</i>. <i>uniformis handfieldi</i> ssp. n. in eastern Canada). <b>Barcode results.</b> A total of 310 North American <i>Lasionycta</i> specimens were submitted for CO1 barcode sequencing. Of these, 208 produced sequences of 600 or more base pairs and 168 were complete (658 bp). No barcode results are available for four species (<i>L</i>. <i>carolynae</i>, <i>L</i>. <i>illima</i>, <i>L</i>. <i>macleani</i>, <i>L</i>. <i>mono</i>) and only incomplete sequences are available for eight more (<i>L</i>. <i>coracina</i>, <i>L</i>. <i>dolosa</i>, <i>L</i>. <i>flanda</i>, <i>L</i>. <i>lagganata</i>, <i>L</i>. <i>phaea</i>, <i>L</i>. <i>sasquatch</i>, <i>L</i>. <i>sierra</i>, <i>L</i>. <i>subfumosa</i>) (all barcode data is presented in the Appendix). The sample set is skewed by the availability of fresh material from the Pacific Northwest, Alberta, and Churchill, Manitoba and a relative paucity of suitable material from most other parts of North America. Most samples are from sub-groups in which variability suggested the possibility of several similar species. <i>Lasionycta uniformis</i>, <i>L</i>. <i>subfuscula</i>, <i>L</i>. <i>perplexa</i>, and <i>L</i>. <i>leucocycla</i> are particularly well sampled. Twenty of the 39 species (51.3 %) with complete or partial sequences had a single CO1 haplotype. Five of these are only known from a single sequence, so the true haplotype diversity of these species is unknown. Eleven species (28.2 %) had two haplotypes, and two species had three (5.1 %). Th ere were no species with four or five haplotypes, but six species (15.4 %) had six to twelve haplotypes (<i>L</i>. <i>leucocycla</i>, <i>L</i>. <i>perplexa</i>, <i>L</i>. <i>promulsa</i>, <i>L</i>. <i>subfuscula</i>, <i>L</i>. <i>taigata</i>, <i>L</i>. <i>uniformis</i>). Eight of 39 species (20.5 %) shared haplotypes (<i>L</i>. <i>skraelingia</i> and <i>L</i>. <i>taigata</i>, <i>L</i>. <i>anthracina</i> and <i>L</i>. <i>flanda</i>, <i>L</i>. <i>poca</i> and <i>L</i>. <i>sasquatch</i>, and <i>L</i>. <i>coracina</i> and <i>L</i>. <i>leucocycla</i>). Th e haplotypes of 8 species differed by only one or two base pairs (<i>L</i>. <i>frigida</i> and <i>L</i>. <i>leucocycla</i>, <i>L</i>. <i>phoca</i> and <i>L</i>. <i>uniformis</i>, <i>L</i>. <i>pulverea</i> and <i>L</i>. <i>sierra</i>, <i>L</i>. <i>staudingeri</i> and <i>L</i>. <i>subfumosa</i>). The neighborjoining trees (Figs 247 and 248) demonstrate that seven species are placed adjacent to or nested amongst multiple similar haplotypes of two polytypic species (<i>L</i>. <i>anthracina</i>, <i>L</i>. <i>flanda</i>, <i>L</i>. <i>frigida</i>, and <i>L</i>. <i>coracina</i> with <i>L</i>. <i>leucocycla</i> <i>;</i> <i>L</i>. <i>phoca</i>, <i>L</i>. <i>gelida</i>, and <i>L</i>. <i>discolor</i> with <i>L</i>. <i>uniformis</i>). Therefore, 17 <i>Lasionycta</i> species (43.6 %) have identical or similar barcodes demonstrating a high frequency of non-discrete haplotype clusters despite the relatively small number of highly polytypic species. Additional sampling of poorly represented species is likely to show an increase in polytypism and haplotype sharing. The CO1 DNA barcodes were helpful for this taxonomic revision in several regards. Above the species level DNA was helpful in defining the generic limits. The DNA data support inclusion of <i>L</i>. <i>secedens</i> and exclusion of <i>Lasionhada proxima</i> and the beach species herein transferred to <i>Psammopolia</i>. Within <i>Lasionycta</i>, distance analysis shows species clustering with relatively large differences between such groups supporting the use of species-groups. Th ese longer DNA tree branches mirror the morphology-based species-group classification described in this paper (Fig. 247). DNA results are less convincing at the sub-group level in the large <i>L</i>. <i>leucocycla</i> species group (Fig. 248). Th e seven sub-groups are comprised of one to ten superficially and structurally similar species. Most of the sub-group members show a tendency to form DNA clusters, but only the <i>L</i>. <i>subfuscula</i> group consisting of a single species lacks overlap with the other sub-groups. Members of the <i>L</i>. <i>staudingeri</i> sub-group are particularly widely dispersed amongst the entire speciesgroup and its species or species pairs differ by over 6 bp from adjacent species. Although this result could be interpreted to mean that the sub-groups are paraphyletic or polyphyletic, it is probably due to presence of shared haplotypes and non-discrete species haplotype clusters. Utility of CO1 DNA was also limited at the species level due to the presence of shared haplotypes, close haplotype similarities, and polytypic species as described above. Despite these shortcomin
description: Published as part of <i>Crabo, Lars &amp; Lafontaine, Donald, 2009, A Revision of Lasionycta Aurivillius (Lepidoptera: Noctuidae) for North America and notes on Eurasian species, with descriptions of 17 new species, 6 new subspecies, a new genus, and two new species of Tricholita Grote, pp. 1-156 in ZooKeys 30 (30)</i> on pages 4-10, DOI: 10.3897/zookeys.30.308, http://zenodo.org/record/576576
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