Revision of the species of Mythimna Ochsenheimer, 1816 (PseudaletiaFranclemont, 1951) (Lepidoptera: Noctuidae: Noctuinae: Leucaniini) occurring in Brazil

ABSTRACT

The genus Mythimna Ochsenheimer, 1816 groups aproximatelly 270 species worldwide. This genus is subdivided into many subgenera and species-groups, with those species occurring in the Neotropical region included in the subgenus Mythimna (Pseudaletia) Franclemont, 1951. Species of this subgenus frequently reach high population levels, causing economic damage to Poaceae cultivars. Hence, it is crucial a well-defined taxonomy allowing rapid and precise identifications. However, the species of this subgenus are cryptic, their recognition requires molecular analyses and in-depth morphological studies, which has often resulted in misidentifications. In Brazil, the occurrence of the following species had been mentioned: Mythimna (P.) adultera (Schaus, 1894), Mythimna (P.) roraimae Franclemont, 1951, Mythimna (P.) sequax Franclemont, 1951, and Mythimna (P.) unipuncta (Haworth, 1809). Yet, until now, we lack a broader knowledge about the geographic distribution and taxonomy of these species. Thus, this study aims to revise the taxonomy of those species based on morphological and molecular (COI) data to provide an updated comprehension of this group in the country. The analysis of 1,664 specimens allowed us to confirm the presence of three species in the country: Mythimna (P.) adultera, Mythimna (P.) sequax, and Mythimna (P.) unipuncta. A lectotype for Mythimna (P.) adultera is designated; a new synonym is proposed: Pseudaletia roraimae syn. nov. of Mythimna (P.) unipuncta; and a new species, Mythimna (P.) celiae sp. nov. is described from Bagé, Rio Grande do Sul, Brazil.

Keywords:
Catalog; Geographic distribution; True armyworm; Morphology; Taxonomy

Introduction

MythimnaOchsenheimer, 1816 (Lepidoptera: Noctuidae: Noctuinae: Leucaniini) is a cosmopolitan genus that comprises approximately 270 species, grouped in subgenera and species-groups according to genitalia characters (Yoshimatsu, 1994, 1995; Yoshimatsu and Hreblay, 1998; Yoshimatsu and Legrain, 2001). This classification has been accepted in the most recent taxonomic studies, though it remains problematic (Hacker et al., 2002; Lafontaine and Schmidt, 2010).

The subgenus Mythimna (Pseudaletia) Franclemont, 1951 is represented by 17 species with ocurrences in Neotropical and Oriental regions and Hawaii islands. It counts with the following distinguishing characters: a distinct and sclerotized valva with ampulla and clasper reduced in size (Hacker et al., 2002). Most of the 17 species belonging to this subgenus are economically important (Hacker et al., 2002), and they frequently reach high population levels, which causes economic damage (Salvadori and Parra, 1990a, 1990b, 1990c; Rizzo and La Rossa, 1991; Capinera, 2006). The following species have been recorded in Brazil: Mythimna (P.) unipuncta (Haworth, 1809), Mythimna (P.) adultera (Schaus, 1894), Mythimna (P.) sequax Franclemont, 1951, and Mythimna (P.) roraimae Franclemont, 1951 (Franclemont, 1951). There are records of the occurrence of Mythimna (P.) punctulata (Blanchard, 1852) in Brazil, Chile, Peru (Artigas, 1994), and Argentina (Artigas, 1994; Pastrana, 2004). However, Franclemont (1951) and Angulo and Weigert (1977) mentioned its occurrence exclusively to Chile.

Mythimna (P.) roraimae is cited only in its original description (Franclemont, 1951) and the world catalog of Noctuidae (Poole, 1989), with distribution restricted to the type locality. On the other hand, Mythimna (P.) sequax is widely distributed in the American continent, from Souther Brazil, Northen of Argentina and Uruguay to United States of América: Florida and Texas (Franclemont, 1951; Heppner, 2007). It is economically important in Brazil, and for this reason, it has been cited hundreds of times in the literature (Gassen, 1984; Madruga et al., 2019). Mythimna (P.) adultera is a common species present from Southern Brazil, Uruguay, Paraguay to Northern part of Argentina. This species is considered of agricultural importance throughout its distribution (Gassen, 1984; Rizzo and La Rossa, 1991; Zerbino, 1991). Finally, Mythimna (P.) unipuncta is known worldwide as a pest (Pond, 1960, 1961; McNeil, 1987; Tavares et al., 1992; Capinera, 2006).

The wide geographic distribution of Mythimna (P.) unipuncta, associated with its economic importance and morphological similarity has led multiple authors to identification mistakes (Biezanko et al., 1949; Biezanko and Bertholdi, 1951; Bertels, 1956; Corseuil, 1958; Costa, 1958; Bertels and Baucke, 1966). The main association came from the works of Hampson (1905) and Draudt (1919), in which several species and forms were considered synonyms of Cirphis unipuncta. Since then, this has been a recurring problem, given that the species of this subgenus are highly similar, most of them distinguishable only through careful examination of their genitalia.

Franclemont (1951) revisited the “unipuncta group” based on morphological evidence, his results being valid to this day. Currently, new tools such as molecular studies have contributed to bringing light to the systematic of the genus (Sutrisno, 2012). But, up till now, we lack a broader knowledge of the geographic distribution and taxonomy of the Brazilian species. Considering their agricultural importance, it is crucial to have these two aspects well-defined, so that the identification of these species is rapid and reliable. This study aims to revise the species of Mythimna (Pseudaletia) occurring in Brazil, by using morphological and molecular (COI) characters to analyze and elucidate its current status and composition in the country.

Material and methods

Collections

The studied specimens belong to the following collections:

AMNH - American Museum of Natural History, New York, U.S.A.

CEUCS - Coleção Entomológica da Universidade de Caxias do Sul, Caxias do Sul, Rio Grande do Sul, Brazil.

CLAM - Coleção Alfred Moser, São Leopoldo, Rio Grande do Sul, Brazil.

DZUP - Coleção Entomológica Padre Jesus Santiago Moure, Departamento de Zoologia, Setor de Ciências Biológicas, Universidade Federal do Paraná. Curitiba, Paraná, Brazil.

CPAC - Coleção Entomológica da Embrapa Cerrados, Planaltina, Distrito Federal, Brazil.

IOC - Instituto Oswaldo Cruz. Rio de Janeiro, Rio de Janeiro, Brazil.

MECB - Museu Entomológico Ceslau Biezanko, Universidade Federal de Pelotas, Pelotas, Rio Grande do Sul, Brazil.

MCTP - Museu de Ciências e Tecnologia da Pontifícia Universidade Católica do Rio Grande do Sul, Porto Alegre, Rio Grande do Sul, Brazil.

MGCL - McGuire Center for Lepidoptera and Biodiversity, University of Florida, Gainesville, Florida, U.S.A.

MNHN - Muséum national d’Histoire naturelle, Paris, France.

MNRJ - Museu Nacional. Universidade Federal do Rio de Janeiro, Rio de Janeiro, Rio de Janeiro, Brazil.

MZUSP - Museu de Zoologia, Universidade de São Paulo, São Paulo, São Paulo, Brazil.

NHMUK - The Natural History Museum, London, United Kingdom.

USNM - National Museum of Natural History, Smithsonian Institution. Washington D.C., U.S.A.

We also collected specimens using light traps model Pennsylvania (Frost, 1957), equipped with fluorescent and black lights, for five nights in each new moon between June, 2015 and May, 2016 in 15 Brazilian localities: Alto Alegre - Roraima; Rio Branco - Acre; Mojuí dos Campos - Pará; Porto Nacional - Tocantins; Petrolina - Pernambuco; Sinop - Mato Grosso; Planaltina - Distrito Federal; Chapadão do Sul and Miranda - Mato Grosso do Sul; Uberaba - Minas Gerais; Alegre and Domingos Martins - Espírito Santo; Londrina - Paraná; Passo Fundo, Vacaria and Bagé - Rio Grande do Sul. After screening and identification, the material was preserved wet (in alcohol 70%) or dry (entomological pins), and part of it was incorporated into the CPAC and DZUP collections.

The nomenclature adopted for venation and genitalia characters is referred to in Fig. 1. The abbreviations used throughout the text are as follows: apl. - applied; biol.- biology; geogr. distr. - geographic distribution; inv.- inventory; morph.- morphology; parasit. - parasitism.

Figure 1
Nomenclature adopted for wing venation and genitalia, based on Mythimna (P.) celiae sp. nov. a, fore and hindwing, scale bar = 10mm; b, male genitalia in ventral view; c, ampulla, digitus, and clasper in inner lateral view of the valva, with a scheme on the right side; d, uncus in posterior view; e, uncus in lateral view; f, fultura inferior (=juxta); g, aedeagus with everted vesica in lateral view; h-i, female genitalia in lateral and ventral views, respectively. Scale bars b-e, g-i= 1mm, f = 0.5mm.

The records of the geographic distribution of the species were taken from the labels of the examined material. The geographic coordinates were obtained through Google Earth, and the geographic distribution map was generated with SampleMappr (Shorthouse, 2010). The examined material corresponds only to specimens deposited in scientific collections. The classification follows Hacker et al. (2002) for genus and subgenus, and Lafontaine and Schmidt (2010) for higher taxonomic categories.

Mythimna (Pseudaletia) unipuncta is redescribed based on images and its catalog is restricted to the main taxonomic works. This was necessary because of the enormous number of taxonomic problems related to this species over time caused by the lack of examination of its type specimens. This problem renders the publications dealing with it untrustworthy, which is not the case for the other species.

Morphological and molecular analyses

A total of 1,664 specimens were examined and length of their forewing measured. For the genitalia character survey, 52 specimens were dissected as in Dias et al. (2017) with nomenclature of Franclemont (1951) and Hacker et al. (2002).

For the molecular analysis, 22 sequences of 658 base pairs of the mitochondrial DNA ‘barcode’ region of the cytochrome c oxidase subunit I (COI) gene were obtained and deposited in Genbank (Sayers et al., 2020). Additionally, were added for comparative analyses 21 sequences mined from Bold Systems (Ratnasingham and Hebert, 2007). Accession numbers and locality information provided in each cladogram. The data was edited in the program BioEdit (Hall, 2011) and analyzed using MEGA11 (Tamura et al., 2021). Evolutionary trees were inferred using the Neighbor-Joining and Maximum Likelihood algorithms in Tamura model 3-parameters with a gamma distribution with Invariant Sites (T92 + G + I), with bootstrap of 1,000 replicates. The initial trees for the heuristic search were automatically obtained by applying the algorithms Neighbor-Joining and BioNJ and by selecting the topology with the highest log-likelihood value. A discrete Gamma distribution was used to model the differences in the evolutionary rate between the sites (5 categories (+ G, parameter = 1,0000)). The trees were drawn in scale, with the average ramification length measured in the number of substitutions per site.

Intra and interspecific evolutionary divergences were estimated with MEGA11, using the same model and bootstrap. Positions containing gaps and missing data were deleted.

Results

Mythimna Ochsenheimer, 1816

MythimnaOchsenheimer, 1816. Schmett. Europ. 4, p. 78.- Walker, 1856. List Spec. Lepid. Insects Coll. Brit. Mus. 9, 72.- Draudt, 1919. Gross-Schmetterlinge Erde 7, 79.- Franclemont, 1951. Proc. Ent. Soc. Wash. 53(2): 60.- Poole, 1989. Lep. Cat. (n.s.) 118, Noctuidae 2, 673; syn.:Philostola Billberg, 1820; Hyperiodes Warren, 1910.- Yoshimatsu, 1994. Bull. Nat. Inst. Agro-Environ. Sci. 11: 81.- Yoshimatsu, 1995. Japn. J. Ent. 63: 235.- Yoshimatsu and Legrain, 2001. Entomol. Sci. 4: 431.- Hacker et al. 2002. Noctuidae Europaeae 4, 168; syn.: Philostola Billberg, 1820; Aletia Hübner, [1821]; Heliophila Hübner, 1822; Barolia Moore, 1886; Hyperiodes Warren, 1910; Hypopteridia Warren, 1912; Conithimna Beck, 1999; Gruathimna Beck, 1999; Foehstia Beck, 1999; Pudothimna Beck, 1999; Ferrathimna Beck, 1999; Allitoria Beck, 1999.- Speidel and Naumann, 2004. Syst. Biodivers. 2(2): 195.- Fibiger and Lafontaine, 2005. Esperiana 11: 13.- Lafontaine and Schmidt, 2010. ZooKeys 40: 91.- Sustrino, 2012. HAYATI J. Biosc., 19(2): 65.

Mythimna (Pseudaletia) Franclemont, 1951

Pseudaletia Franclemont, 1951. Proc. Ent. Soc. Wash. 53(2): 63.- Franclemont and Todd, 1983. Check List Lep. Amer. N. Mexico, 150.- Poole, 1989. Lep. Cat. (n.s.) 118, Noctuidae 2, 843.

Mythimna (Pseudaletia); Zerbino, 1984. Investig. Agron. (Montevideo) 5: 17.- Terra and Zerbino, 1986. Investig. Agron. (Montevideo) 6: 49.- Holloway, 1989. Moths of Borneo, Part 12, Malay. Nat. J. 42(2-3): 57.- Yoshimatsu, 1994. Bull. Nat. Inst. Agro-Environ. Sci. 11: 81.- Hacker et al., 2002. Noctuidae Europaeae 4, 178.- Barbut and Lalanne-Cassou, 2010. L’Ent. 66(3): 117.- Sutrisno, 2012. HAYATI J. Biosc. 19(2): 65.

Systematic history

Ochsenheimer described Mythimna in 1816 based on Phalaena turca Linnaeus, 1761, as the type species. Posteriorly, Franclemont (1951) described the genus Pseudaletia with Noctua unipuncta Haworth, 1809 as the type species. Franclemont and Todd (1983) and Poole (1989) maintained the status of genus for Pseudaletia. Zerbino (1984) and Terra and Zerbino (1986) cited Pseudaletia as a synonym of Mythimna. However, they mentioned that no taxonomic justification could be found to include the South American species in the latter. Holloway (1989) cited Pseudaletia as subgenus of Mythimna. Yoshimatsu (1994) revised the species from Japan and Taiwan, finding genus Mythimna monophyletic and confirming the presence of seven subgenera, Pseudaletia among them. Finally, Hacker et al. (2002) followed the same hypothesis, citing the Neotropical species in the subgenus Mythimna (Pseudaletia).

Morphological characterization

The species of Mythimna are predominantly beige, a pattern that is generally mixed with dark scales. Many species are superficially similar, and the distinction often requires examination of the genitalia. As a standard, male and female genitalia differ in general appearance, with emphasis on the valva and its structures in males and the structure of the bursa copulatrix in females (Franclemont, 1951; Hacker et al., 2002).

In the species of Mythimna (Pseudaletia), the forewings are narrower and have a sharper apex when compared to the wing morphology of the other subgenera. Its genitalia has the following synapomorphies: male genitalia with the harpe-ampulla complex reduced and modified, cucullus expanded, corona strong and extended, vesica usually with a basal diverticulum bearing a reduced, terminal cornutus. Female genitalia with appendix bursae deriving from the proximal part of ductus bursae (Hacker et al., 2002).

Morphological and molecular analyses have agreed with the presence of four species in Brazil: Mythimna (P.) unipuncta (Haworth, 1809), Mythimna (P.) adultera (Schaus, 1894), Mythimna (P.) sequax Franclemont, 1951, and Mythimna (P.) celiae sp. nov. (Figs. 1, 2, 3, 4; Table 1).

Figure 2
Geographic distribution of Mythimna (Pseudaletia) in Brazil.

Figure 3
Evolutionary relationships of the taxa, using the Neighbor-Joining method. The optimal tree with the sum of the branch lengths = 0,14304783 is shown, bootstrap values are presented next to each ramification.

Figure 4
Molecular phylogenetic analysis through the method of maximum likelihood (original tree). The tree with the highest log probability (-1360.63) is shown. The percentage of trees in which the grouped associated taxa is shown beside the ramifications.

Table 1
Estimates of evolutionary divergence over pairs of sequences between groups and within groups. The standard error estimates are in grey. Sample number (n).

	N	D.	S.E.		M. (P.) celiae sp. nov.	M. (P.) sequax	M. (P.) adultera	M. (P.) unipuncta	M. oxygala	M. yukonensis
					S.E.
M. (P.) celiae sp. nov.	02	0.001	-	Between species		0.008	0.007	0.005	0.013	0.013
M. (P.) sequax	23	0.001	0.001		0.041		0.007	0.008	0.014	0.014
M. (P.) adultera	06	0.001	0.001		0.028	0.028		0.007	0.014	0.013
M. (P.) unipuncta	08	0.000	-		0.017	0.040	0.027		0.014	0.014
M. oxygala	02	0.001	-		0.082	0.093	0.091	0.091		0.005
M. yukonensis	02	0.000	-		0.083	0.093	0.086	0.089	0.015

Mythimna (Pseudaletia) unipuncta ( Haworth, 1809 ) (Figs. 5, 6, 7)

Figure 5
Mythimna (Pseudaletia) unipuncta. a - male lectotype in dorsal view. Scale bar = 1cm. b: male genitalia in posterior view with aedeagus in lateral view. Specimen from NHMUK 010354729. There is no scale in the genitalia.

Figure 6
Mythimna (Pseudaletia) unipuncta. a: female in dorsal view. Scale bar = 1cm. b: female genitalia in ventral view. Specimen identified as Leucania antica at the NHMUK 010915614. There is no scale of the genitalia.

Figure 7
Mythimna (Pseudaletia) unipuncta. a: male in dorsal view. b: male genitalia in posterior view with aedeagus in lateral view. Holotype of Pseudaletia roraimae at the AMNH 29780. There are no scales.

Noctua unipuncta Haworth, 1809. Lepid. Brit. 2, 174.- Hampson, 1905. Cat. Lep. Phalaenae 5, 547.

Leucania impuncta [sic]; Stephens, 1829. Illustr. Brit. Entomol. 3, 80.

Leucania unipuncta; Stephens, 1829. Illustr. Brit. Entomol. 3, 338.- Butler, 1890. Trans. Ent. Soc. Lond. 1890(4), 661.- Hampson, 1894. Fauna Brit. India 2, 275; syn.: Leucania extranea, Leucania antica, Leucania convecta, Leucania separata, Leucania adusta, Leucania consimilis, Leucania trifolii, Leucania saccharivora.- McDunnough, 1938. Mem. South. Calif. Acad. Sci. 1, 77; syn.: Leucania extranea.- Koutsaftikis, 1974. Ann. Goulandris Museum. 2, 96.- Poole, 1989. Lep. Cat. (n.s.) 118, Noctuidae 2, 844; syn.: Leucania extraneaGuenée, 1852, Leucania anticaWalker, 1856, Leucania unipuncta quechua Franclemont, 1951, Leucania unipuncta tseki Koutsaftikis, 1974.

Leucania extraneaGuenée, 1852. Hist. Nat. Ins., Spec. Gén. Lépid. 5, 77.- Walker, 1856. List Spec. Lepid. Insects Coll. Brit. Mus. 9, 93.- Hampson, 1894. Fauna Brit. India 2, 275.- Hampson, 1905. Cat. Lep. Phalaenae 6, 547.- Franclemont, 1951. Proc. Ent. Soc. Wash. 53, 65, lectotype, without sex or locality, deposited in USNM.- Viette, 1951. Bull. mens. Soc. Linnéenne Lyon 20(7), 159; lectotype female, Montevideo, coll. Feisthamel, MNHN.

Leucania anticaWalker, 1856. List Spec. Lepid. Insects Coll. Brit. Mus. 9, 100.- Hampson, 1894. Fauna Brit. India 2, 275.- Hampson, 1905. Cat. Lep. Phalaenae 5, 547.

Leucania unipuncta var. antica; Butler, 1890. Trans. Ent. Soc. Lond. 1890(4), 661.

Leucania unipuncta var. extranea; Butler, 1890. Trans. Ent. Soc. Lond. 1890(4), 662.

Cirphis unipuncta; Hampson, 1905. Cat. Lep. Phalaenae 5, 547; syn.: Spaelotis punctulata, Leucania extranea, Leucania antica, Leucania convecta, Leucania separata, Leucania consimilis, Leucania trifolii, Leucania saccharivora, Leucania adultera.- Draudt, 1919. Gross-Schmetterlinge Erde 12, 167; syn.: Leucania extranea, Leucania antica, Leucania trifolii, Leucania adultera.-Specht and Corseuil, 1996. Biociências 4(2), 146.

Leucania unpuncta [sic]; Franclemont, 1951. Proc. Ent. Soc. Wash. 53(2), 64.

Pseudaletia unipuncta; Franclemont, 1951. Proc. Ent. Soc. Wash. 53(2), 65, figs 10, 10a, 11, 11a (male gen.), 33 (female gen.); syn.: Leucania extranea.- Godfrey, 1972. Tech. Bull. 1450, 86.- Nye, 1975. Gen. Nam. Moth World 1, 410; lectotype male, Great Britain: [England] Anglia (BMNH).- Franclemont and Todd, 1983. Check List Lep. Amer. N. Mexico, 150.- Poole, 1989. Lep. Cat. (n.s.). 118, Noctuidae 2, 844; syn.: Leucania extranea, Leucania antica, Pseudaletia unipuncta quechua, Leucania unipuncta tseki.

Pseudaletia unipuncta antica; Franclemont, 1951. Proc. Ent. Soc. Wash. 53(2), 66, figs 12, 12a (male gen.), 34 (female gen.).

Pseudaletia unipuncta quechua Franclemont, 1951. Proc. Ent. Soc. Wash. 53(2), 67, figs 13, 13a (male gen.).

Pseudaletia roraimae Franclemont, 1951. Proc. Ent. Soc. Wash. 53(2), 68, figs 15, 15a (male gen.).- Poole, 1989. Lep. Cat. (n.s.). 118, Noctuidae 2, 844. New synonymy.

Leucania unipuncta tseki Koutsaftikis, 1974. Ann. Goulandris Museum 2, 96.

Mythimna (Pseudaletia) roraimae; Hacker et al., 2002. Noctuidae Europaeae 4, 168.

Mythimna (Pseudaletia) unipuncta; Hacker et al., 2002. Noctuidae Europaeae 4, 179.

Mythimna unipuncta; Lafontaine and Schmidt, 2010. Zookeys 40, 91.

Systematic history

Haworth (1809) described Noctua unipuncta, and later came the descriptions of Leucania extranea Guené, 1852 and Leucania anticaWalker, 1856. Stephens (1829) transferred N. unipuncta to Leucania.Butler (1890) mentions L. antica and L. extranea as varieties of L. unipuncta.Hampson (1905) and Draudt (1919) transferred it to Cirphis Walker, 1865 and treat these variations, var. saccharivora Butler, 1882, var. trifoliiButler, 1882, var. convecta Walker, 1857 and var. separata Walker, 1865, as synonyms of Cirphis unipuncta.Franclemont (1951) in a thorough study that included genitalia morphology, revealed the incorrect classification in Cirphis, given that the American species do not correspond to the description of this genus. Thus, Franclemont (1951) described the genus Pseudaletia, designating Noctua unipuncta as its type species; he also revalidated Leucania antica as a subspecies of Pseudaletia unipuncta and described Pseudaletia unipuncta quechua and Pseudaletia roraimae. Leucania unipuncta tseki was described by Koutsaftikis in 1974; this species and the above-mentioned subspecies were treated as synonyms of Pseudaletia unipuncta by Poole (1989), while Hacker et al. (2002) cited unipuncta in Mythimna (Pseudaletia), and Lafontaine and Schmidt (2010) considered it in Mythimna.

Type material

Noctua unipuncta Haworth, 1809. Described from Anglia (England) without any mention of the number of specimens and sex. Lectotype designated by Nye (1975) and deposited at the NHMUK, with the following labels: /NHMUK010354729/ Stainton Coll. 1893-134./ Type/ Agrotidae genitalia slide No. 896 ♂/ unipuncta [illegible]/.

Leucania extraneaGuenée, 1852. Described from North America, Brazil, Colombia, etc. (several collections), Java [Oceania], East Indies (Collections of the India Company), and Nouvelle Hollande [New York] (M.N. [MNHN]) without any mention of the number of specimens or their sex. Two lectotypes have been designated for this taxon, one by Franclemont (1951, published on April), without sex or locality, with the number 60993 and deposited at the USNM. The other was designated by Viette (1951, published on September); it is a female from Montevideo [Uruguay], from Feisthamel collection, deposited at the MNHN. Thus, the first designation is valid, confirming what had already been said by Poole (1989). The lectotype designated by Franclemont (1951) was not found in the above-mentioned collection (Paul Goldstein, curator of the collection, personal communication, 2018).

Leucania anticaWalker, 1856. Described from the West Coast of the United States, without any mention of the number of specimens or their sex (NHMUK). Hampson (1905) cited 4 ♀ [syn]types from Venezuela deposited at the NHMUK; however, these specimens cannot belong to the type series, since they are not from the type locality. The series of syntypes from the West Coast of the US has not been found, rendering a lectotype designation impossible. In case its existence cannot be attested, a neotype should be designated.

Pseudaletia unipuncta quechua Franclemont, 1951. Subspecies described based on the male holotype from Incachaca, Cochabamba, Bolívia, a male from Napo, Equador, another male from Env[iron]. d’Ambato, Equador, two males and four females from Aqualani, Carabaya, Peru. Male holotype collected by J. Steinbach [leg], Type Nº. 60994, deposited at the USNM with the following labels: /Incachaca Cochabamba Bolivia JSteinbach/ Collection WmSchaus/ ♂ Genitalia Slide: USNM 1182 J. G. Franclemont/ HOLOTYPE Pseudaletia unipuncta quechua J. G. Franclemont/ USNMENT (QRcode) 00973212/.

Pseudaletia roraimae Franclemont, 1951. Species described based on the male holotype from Monte Roraima, [Uiramutã], Roraima, Brazil, deposited at the AMNH, with the following labels: /Mt. Roraima, Brazil Alt. Summit 8860 No. 1927/ Acc. 29780 Tate No./ ♂ Genitalia Slide A-69 J.G. Franclemont/ HOLOTYPE Pseudaletia roraimae J.G. Franclemont/. /NO. A-69 TYPE Pseudaletia roraimae Braz[il]. Franc[lemont]. DATE 19 jan 1951 J.G. FRANCLEMONT/.

Leucania unipuncta tseki Koutsaftikis, 1974. Subspecies described based on the male [holotype] from de Simi, Greece, deposited at the Goulandris Museum, Greece. The [holotype] could not be found (Maria Dimaki, curator of the collection, personal communication, 2018).

Diagnosis

Both sexes with beige or orange-beige forewings, mixed with dark scales. In males, cucullus with a straight dorsal margin, usually ending in a distal process; vesica with a basal diverticulum, bearing more than four cornuti, all closely positioned, the distalmost cornutus being thinner and longer.

Description

MALE. Head: Beige or orange-beige, mixed with dark scales of different hues. Eyes brown and pilose. Antenna beige dorsally and brown ventrally, both surfaces bearing microsetae. Thorax: Ground color as head; patagium with dark subapical line. Forewing: length 2cm (n=1). Ground color beige to orange-beige, mixed with dark scales of different hues, postmedial line as dark spots over the veins and terminal line as dark spots between veins, other transversal lines absent; with an oblique line of dark scales from the apex to M₂, rarely with an anterior area lighter than ground color; a dark patch with a spot of central white scales next to the bases of M₂ and M₃; dark patch over cubital vein absent in most specimens; orbicular and reniform spots circular, inconspicuous, orange with smooth darker center; subterminal area of the wing with white or light beige scales over the veins; fringe beige with dark medial and terminal lines. Hindwing: pale, with a marginal brown shading toward the base; fringe pale. Abdomen: Dorsally greyish-beige; ventral view as head, with two black transversal lines subventral. Genitalia: Tegumen and uncus fused, the former wider than base of uncus, ventral arms robust in posterior view, covered with setae near the articulation with the dorsal arm of saccus. Saccus round and bent toward the ventral arm of the tegumen, with an irregular distal portion, twisted toward its inner surface. Uncus elongated, flat and bent ventrally, widened on medial area, apical half abruptally narrowed to 1/5-1/6 of its width, covered with setae dorsally. Valva subtriangular; sacculus projected ventrally, forming an oval lobe; cucullus elongated, with a straight dorsal margin and usually ending in a distal process; distal portion lobate, corona present, with multiple rows of spines; clasper short; ampulla distinct, as a small ear-shaped dorsal projection; digitus slim, as long as clasper. Fultura inferior (=juxta) subtriangular, with approximately one third the length of the ventral arm of the tegumen and thrice as long as its smallest width, distal portion bifid. Aedeagus pipe-shaped; vesica tubular and elongate, with a continuous band of cornuti from posterior the diverticulum to subapical area, basal diverticulum elongate, with over four closely arranged cornuti, the distalmost thinner and longer than the others.

FEMALE. Similar to male. Forewing length 1.8cm (n=1). Abdomen: Genitalia almost totally sclerotized. Eighth tergite straight and narrow. Papilla analis subtriangular covered with setae. Lamella antevaginalis subrectangular. Ductus bursae sclerotized and with a wide fold at the last portion, which leads to appendix bursae; corpus bursae membranous and round; appendix bursae more than twice as long as ductus bursae, with its same width, sclerotized basally and with membranous apex. Ductus seminalis originating on appendix bursae apex.

Geographic distribution

Northern U.S.A. to North Brazil (Roraima), South and Southwest Europe, Western Asia, and Central Africa (Franclemont, 1951; Capinera, 2006).

Host plants

Native and cultivated grass (Poaceae) such as barley, corn, rye, oat, rice, sugar cane, sorghum, and wheat, but there are cases of damage to other cultures such as carrot, bean, cabbage, celery, cucumber, sweet potato, pepper, and watermelon (Capinera, 2006).

Discussion

Mythimna (P.) unipuncta, restricted to the extreme North of Brazil, is similar to Mythimna (P.) sequax, widely distributed in the country (Madruga et al., 2019). However, both species are easily distinguished from each other through the exam of their genitalia, especially male genitalia. The former species differs in the basal diverticulum, which bears a sequence with more than four closely arranged cornuti, while in Mythimna. (P.) sequax it usually has one cornutus, rarely more, up to four. Yet, misidentifications are common in Brazilian collections, and all the revised records are of Mythimna (P.) sequax or, still, of Mythimna (P.) celiae sp. nov.

Pseudaletia roraimaesyn. nov. has the male genitalia very similar to that of Mythimna (P.) unipuncta, and according to Franclemont (1951), the difference between them is represented by the robustness of the genitalia of P. roraimae. However, considering that the species was described based on a single male from Monte Roraima, [Uiramutã], Brazil, and no other specimen could be found in either the examined collections or the material collected, including that collected close to the type locality (Alto Alegre - Roraima), we opted for synonymizing it under Mythimna (P.) unipuncta.

The original description of Leucania extraneaGuenée, 1852 mentions that it occurs in Brazil; however, since this material could not be found and given that there was no mention to the specific collecting site, the distribuition of Mythimna (P.) unipuncta in the country is restricted to the information of Mythimna (P.) roraimae.

Material examined

Images of the wings and genitalia of the lectotype of Mythimna (P.) unipuncta, of the “syntype” of Leucania antica, with the following labels: /NHMUK 010915614/ Type/ Agrotidae genitalia slide No 852/ Venezuela 47-9./ 52. Leucania antica./ and of the holotype of Pseudaletia roraimae, in addition to the illustrations in Franclemont (1951) and molecular data from Bold Systems.

Mythimna (Pseudaletia) adultera ( Schaus, 1894 ) (Figs. 8, 9)

Figure 8
Mythimna (Pseudaletia) adultera. a-b, male in dorsal and ventral views; c-d, female in dorsal and ventral views. Specimen deposited at the DZUP. Scale bar = 1cm.

Figure 9
Mythimna (Pseudaletia) adultera. a, male genitalia in left lateral view; b, saccus in dorsal view; c: detail of the ampulla, digitus and clasper of the right valva in inner view; d, fultura inferior in dorsal view; e, aedeagus in left lateral view with everted vesica; f, female genitalia in left lateral view. Scale bars = 1mm.

Leucania adultera Schaus, 1894. Trans. Amer. Ent. Soc. 21, 232.- Hampson, 1905. Cat. Lep. Phalaenae 5, 547.- Specht and Corseuil, 1996. Biociências 4(2), 146; inv., syst.

Cirphis unipuncta [misidentifications]; Biezanko et al., 1949. Agros 2(3), 196; inv., host plant.- Biezanko and Bertholdi, 1951. Agron. 10(4), 244; syn.: Cirphis extranea, Cirphis antica, Cirphis trifolii; inv., host plant.- Bertels, 1956. Ent. Agric. Sul-Brasileira, 346; fig. 193 (egg, larva, pupa); morph., host plant.- Bertels and Baucke, 1966. Pesq. agropec. bras. 1, 30; inv., host plant.- Specht and Corseuil, 1996. Biociências 4(2), 146; inv., syst.

Pseudaletia adultera; Franclemont, 1951. Proc. Ent. Soc. Wash. 53(2), 69, figs 16, 16a (male gen.), 36 (female gen.); syst.- Treat, 1966. Jour. N. Y. Ent. Soc. 74(3), 157 parasit.- Silva et al., 1968. Quarto Cat. Ins. Viv. Plantas Brasil 2(1), 231; apl., host plant, inv., parasit.- Biezanko and Ruffinelli, 1971. Centro Invest. Sanid. Veg. Montevideo, Ser. Zool. Agric., Publ. Técn. 2, 16-17; apl., host plant, inv., parasit.-Casella and Moratoria, 1971. Rev. peruan. Ent. 14(2), 259, figs 1-4 (pupa), 5-8 (larva); host plant.- Carvalho et al., 1971. Rev. Centro Ciências Rurais 1(3), 17.- Morey, 1971. Rev. peruan. Ent. 14(2), 263.- Biezanko et al., 1974. Rev. Centro Ciências Rurais 4(2), 118; host plant.- Corseuil and Cruz, 1975. Rev. Faculdade de Agronomia UFRGS 1(1), 19; inv.- Tarragó et al., 1975. Rev. Centro de Ciências Rurais5(2), 125; inv.- Link, 1977. Rev. Centro de Ciências Rurais 7(4), 331; inv.- Chiang, 1978. Ann. Rev. Ent. 23, 105; apl., host plant.- Zerbino et al., 1983. Investig. Agron. 4, 20; apl.- Poole, 1989. Lep. Cat. (n.s.) 118, Noctuidae 2, 843.- Rizzo and La Rossa, 1991. Rev. Fac. Agron. 12(1), 39, figs 2 (male and female, pupa), 3 (male gen.); biol.- Zerbino, 1991. INIA, Sér. Téc. 9, 1, figs 1 (larva), 2 (adult); biol., apl.- Igarzábal et al., 1994. Gayana Zool. 58(2), 108, fig. 81 (larva); key.- Zerbino, 1994. INIA, Sér. Téc. 47, 1; biol.- Specht and Corseuil, 1996. Biociências 4(2), 146; inv., syst.- Matrangolo et al., 1997. Pesq. agropec. bras. 32(8), 777; host plant.- Ferreira, 1998. Manual de Identificação de Pragas do Arroz, EMBRAPA, 80, figs 74 (adult), 76 (larva); biol.- Specht and Corseuil, 2002b. Rev. Bras. Zool. 19(Supl.1), 286; inv.- Specht et al., 2004. R. Bras. Agrociências 10(4), 398; inv., host plant.- Barnes and Convey, 2005. Antartic Science 17(3), 307; geogr. distr.- Specht et al., 2005. Rev. Bras. Ent. 49(1), 136; inv.- Fritz et al., 2008. Oecol. Bras. 12(4), 723; apl., host plant.- Hernandes et al., 2011. Zootaxa 2936, 11; parasit.- Fritz et al., 2013. Intern. Jour. Trop. Ins. Sci. 33(3), 179; apl.

Mythimna adultera; Zerbino, 1984. Investig. Agron. 5, 17; biol.- Terra and Zerbino, 1986. Investig. Agron. 6, 49; biol.- Fernandes et al., 2014. EntomoBrasilis 7(2), 128; parasit.

Psendaletia [sic] adultera; Specht and Corseuil, 1996. Biociências 4(2), 146; inv., syst.

Mythimna (Pseudaletia) adultera; Hacker et al., 2002. Noctuidae Europaeae 4, 168.- Soria and DeGrande, 2011. Rev. Bras. Milho e Sorgo 10(2), 103; inv.

Systematic history

Leucania adultera was transferred to Pseudaletia by Franclemont (1951), and so far, it has been most often cited in this combination. Zerbino (1984) and Terra and Zerbino (1986) mention it in Mythimna.

Type material

Leucania adultera Schaus, 1894. Species described without any mention of the number of specimens or their sex. There is a male syntype from Castro, Paraná, Brazil, deposited at the USNM, with the following labels: /Castro. Paraná./ Collection WmSchaus/ Leucania unipuncta Haw./ Leucania adultera Type, Schs./ ♂ Genitalia Slide: USNM 1178 J. G. Franclemont/ Type No. 11175 U.S.N.M./. In order to guarantee the correct identification of the species, this specimen is here designated as the lectotype, and the following labels will be added to it: /Lectotypus/ Lectotypus Leucania adultera Schaus, 1894. Madruga, Specht, San Blas, Mielke & Casagrande det. 2022/. These labels will be sent to the Lepidoptera curator of the referred museum.

Diagnosis

Both sexes with greyish-beige forewings, usually mixed with dark scales and with a patch of dark scales over the cubital vein. In males, cucullus with a protuberant dorsal margin, ending in a truncate ventral process, ampulla absent (as a tiny ear-shaped dorsal projection on some specimes), digitus slim, half as long as clasper; vesica with basal diverticulum absent.

Description

MALE. Head: Greyish-beige, usually mixed with dark scales of different hues. Eyes brown, pilose. Antenna filiform, triangular in cross section, covered dorsally with beige scales, ventrally naked, with tiny chemoreceptive trichoid sensilla, more concentrated in the central part of the antennomer, and with longer lateral chemoreceptive trichoid sensilla, almost as long as antennomer width. Thorax: Ground color as head; patagium with dark subapical line. Forewing: length 1.6-1.8cm (n=29). Ground color greyish-beige, thickly mixed with dark scales of different hues; postmedial line as dark spots over the veins and terminal line as dark spots between veins, other transversal lines absent; with an oblique line of dark scales from the apex to M₂, usually with an anterior area lighter than ground color; a dark patch with a spot of white central scales near the base of M₂ and M₃, undifferentiable in some specimens; and a dark long patch usually covering cubital vein; orbicular and reniform spots circular, inconspicuous, orange with smooth darker center; subterminal area of the wing with white or light beige scales over the veins; fringe beige with dark medial and terminal lines. Hindwing pale, with a marginal brown shading toward the base, fringe pale. Abdomen: Greyish-beige in dorsal view; ventral view as head, with two black transversal lines subventral. Genitalia: Tegumen and uncus fused, the former wider the than base of uncus; ventral arms robust in posterior view, covered with setae near the articulation with the dorsal arms of saccus. Saccus round and bent towards the ventral arm of tegumen, irregular distally, twisted toward the inner surface. Uncus elongated, flat and bent ventrally, widened on medial area, apical half abruptally narrowed to 1/6-1/7 of its width, covered with setae dorsally.Valva subtriangular; saccullus projected ventrally, forming an oval lobe; cucullus elongated, with dorsal margin protuberant and ending in a wide, single, truncated distal process, distal portion lobate, corona present, with multiple rows of spines; clasper short; ampulla absent on most specimens, as a tiny ear-shaped dorsal projection rarely; digitus slim, half as long as clasper. Fultura inferior (=juxta) subrectangular, approximately half the length of the ventral arms of tegumen and twice as long as its smallest width, distal portion bifid. Aedeagus pipe-shaped; vesica tubular, wide in basal third then abruptally narrowed to half its width and slightly widened apically, with a continuous band of cornuti, beginning in the right basal area, turning to the left dorsally, almost completing a spin, then in an almost straight line to the subapical area, subbasal cornuti larger than the others, basal diverticulum absent.

FEMALE. Similar to male, but antenna with tiny chemoreceptive trichoid sensilla only, longer lateral chemoreceptive trichoid absent and forewing less densely mixed with dark scales, which gives it a more “clean” and smooth appareance. Forewing length 1.3-2cm (n=35). Abdomen: Genitalia almost entirely sclerotized. Eighth tergite narrow and straight. Papilla analis subtriangular and covered with setae. Lamella antevaginalis subrectangular. Ductus bursae sclerotized, with a wide fold at its final portion leading to appendix bursae; corpus bursae membranous and round; appendix bursae more than twice the length of ductus bursae and half its width, sclerotized basally and with membranous apex. Ductus seminalis originating on appendix bursae apex.

Geographic distribution

Paraguay, Uruguay, Northern Argentina (Franclemont, 1951), and central Argentina (San Blas et al., 2021). Brazil: Paraná: Tibagi, Castro, Ponta Grossa, Quatro Barras, and Morretes; Santa Catarina: Seara; Rio Grande do Sul: Barracão, Rondinha, Alegrete, Santa Maria (Carvalho et al., 1971; Specht et al., 2004; Specht et al., 2005), Iraí, Passo Fundo, Vacaria, São José dos Ausentes, Bento Gonçalves, Salvador do Sul, Cachoeira do Sul, Encruzilhada do Sul, Tavares, Bagé, and Pelotas.

Host plants

Native and cultivated grasses such as oat, barley, canary grass, annual ryegrass, rye, wheat, corn, rice, sword grass and other grasses (Biezanko and Ruffinelli, 1971; Casella and Moratoria, 1971; Biezanko et al., 1974; Chiang, 1978; Matrangolo et al., 1997; Pastrana, 2004; Specht et al., 2004; Fritz et al., 2008).

Discussion

Mythimna (P.) adultera is usually smaller and greyer than the other species. The patch of dark scales over the cubital vein on the dorsal forewing is a determining character in both sexes, but it is not always clear, especially in females. The male genitalia is the most characteristic structure that allows the undoubted differentiation among all other South American species of the genus, in which the dorsal margin of cucullus is protuberant, the uncus and fultura inferior wider than in the other species and the vesica is tubular, without diverticulum, being the only species lacking this structure.

Material examined

BRAZIL - Paraná: Morretes – Parque Estadual Pico Marumbi, Pedra Lascada, 25°26'18,4”S, 48°55'09,6”W, 510m, 1 male, 2 females, 29-IX-3-X-2013, Silva leg., (MZUSP); Quatro Barras - Banhado, 800m, 1 female, 30-X-1970, Becker & Laroca leg., DZ 37.872 (DZUP); Tibagi – Parque Estadual do Guartelá, 24°33'44”S, 50°15'33”W, 950-1000m, 1 male, 25-27-VII-2017, Dantas, Dias, Dolibaina & Queiroz-S[antos] leg., DZ 37.871 (DZUP). - Santa Catarina: Seara - Nova Teutônia, 27°11’S, 52°23’W, 300-500m, 1 male, V-1976, Fritz Plaumann leg., DZ 37.895 (DZUP). - Rio Grande do Sul: Bagé - 31°21'4.94”S, 54°1'12.51”W, 232m, 1 female, 5-XII-2015, R. N. Systi, J. U. P. Corrêa, M. A. P. da Silva & Harry Ebert leg., (CPAC), 1 female, 3 males, 2-II-2016, R. N. Systi, J. U. P. Corrêa, M. A. P. da Silva & Harry Ebert leg., DZ 39.978 (DZUP, CPAC), 2 females, 4-II-2016, R. N. Systi, J. U. P. Corrêa, M. A. P. da Silva & Harry Ebert leg. (CPAC), 2 males, 6-II-2016, R. N. Systi, J. U. P. Corrêa, M. A. P. da Silva & Harry Ebert leg., DZ 39.979 (DZUP, CPAC), 1 female, 2 males, 8-III-2016, R. N. Systi, J. U. P. Corrêa, M. A. P. da Silva & Harry Ebert leg., DZ 39.976 (DZUP, CPAC), 1 male, 1 female, 14-IV-2016, R. N. Systi, J. U. P. Corrêa, M. A. P. da Silva & Harry Ebert leg. (CPAC), 1 male, 4-V-2016, R. N. Systi, J. U. P. Corrêa, M. A. P. da Silva & Harry Ebert leg., DZ 37.896 (DZUP), 1 female, 25-XI-2016, R. N. Systi, J. U. P. Corrêa, M. A. P. da Silva & Harry Ebert leg. (CPAC), 1 male, 6-XII-2015, R. N. Systi, J. U. P. Corrêa, M. A. P. da Silva & Harry Ebert leg., DZ 37.898 (DZUP), 31°18'57.08”S, 53°59'52.91”W, 242m, 1 male, 7-X-2015, R. N. Systi, J. U. P. Corrêa, M. A. P. da Silva & Harry Ebert leg., DZ 37.893 (DZUP), 1 female, 2 males, 6-XII-2015, R. N. Systi, J. U. P. Corrêa, M. A. P. da Silva & Harry Ebert leg. (CPAC), 1 male, 7-XII-2015, R. N. Systi, J. U. P. Corrêa, M. A. P. da Silva & Harry Ebert leg., DZ 37.870 (DZUP), 2 females, 11-III-2016, R. N. Systi, J. U. P. Corrêa, M. A. P. da Silva & Harry Ebert leg., DZ 39.977 (DZUP, CPAC); Bento Gonçalves – 1 male, 19-IV-2004, Specht leg. (CEUCS); Cachoeira do Sul – 1 female, 21-V-1998, Specht leg., 11558 (MCTP); Encruzilhada do Sul - 400-500m, 2 females, 2-3-XI-2000, A. Moser leg. (CLAM); Iraí - 1 female, 20-X-1998, J. A. Teston leg., 11562 (MCTP); Passo Fundo - 28°13'50.67”S, 52°24'17.04”W, 671m, 1 male, 11-X-2015, P. R. V. S. Pereira leg., DZ 37.900 (DZUP), 1 male, 8-XI-2015, P. R. V. S. Pereira leg. (CPAC), 1 female, 9-XI-2015, P. R. V. S. Pereira leg. (CPAC), 28°13'35.88”S, 52°24'13.05”W, 682m, 2 females, 9-X-2015, P. R. V. S. Pereira leg. (CPAC), DZ 37.892 (DZUP), 1 female, 11-X-2015, P. R. V. S. Pereira leg., DZ 37.877 (DZUP), 1 male, 14-II-2016, P. R. V. S. Pereira leg., DZ 37.901 (DZUP), 2 females, 8-III-2016, P. R. V. S. Pereira leg. (CPAC), 1 female, 1 male, 10-III-2016, P. R. V. S. Pereira leg. (CPAC), 1 male, 11-III-2016, P. R. V. S. Pereira leg. (CPAC), 1 female 4-IV-2016, P. R. V. S. Pereira leg., DZ 37.869 (DZUP), 1 female, 1-VI-2016, P. R. V. S. Pereira leg., DZ 37.897 (DZUP); Pelotas – 1 female, 21-I-1951, C. Biezanko leg. (MECB), 1 female, 11-IX-1951, C. Biezanko leg. (MECB), 2 females, I-1954, ex. coll. F. Justus, DZ 37.874, DZ 37.873 (DZUP), 1 male, 6-XII-1957, J. Lucia Mantovani & C. Biezanko leg. (MECB); Salvador do Sul - 1 female, 14-IX-1995, A. Specht leg., 4494 (MCTP), 2 males, 3 females, 4-XI-1997, A. Specht, 11560, 11569, 11563, 11568, 11564 (MCTP); São José dos Ausentes - Silveira, 1200m, 1 male, 28-31-I-2000, A. Moser leg., 1769 (CLAM), Fazenda Potreirinhos, 1200m, 1 male, 5-7-XI-2004, A. Moser leg. (CLAM); Tavares - 1female, 29-X-2010, ex larvae (CEUCS); Vacaria - 1 female, 19-III-2009, A. C. Formentini leg (CEUCS).

Mythimna (Pseudaletia) sequax Franclemont, 1951 (Figs. 10, 11)

Figure 10
Mythimna (P.) sequax. a-b: male (DZ 39.925) in dorsal and ventral views; c-d: female (DZ 39.926) in dorsal and ventral views. Specimen deposited at the DZUP. Scale bar = 1cm.

Figure 11
Mythimna (P.) sequax. a, male genitalia in left lateral view; b, aedeagus in left lateral view with the vesica everted; c, saccus in dorsal view; d, detail of the ampulla, digitus and clasper of right valva in inner view; e, fultura inferior in dorsal view; f, female genitalia in left lateral view. Scale bars = 1mm.

Pseudaletia sequax Franclemont, 1951. Proc. Ent. Soc. Wash. 53(2), 70, figs 18, 18a (male gen.) e 38 (female gen.).- Biezanko and Ruffinelli, 1971. Centro Invest. Sanid. Veg. Montevidéo, Ser. Zool. Agric., Publ. Técn. 2, 17; apl., host plant., inv., parasit.- Carvalho et al., 1971. Rev. Centro Ciências Rurais 1(3), 17; inv.- Biezanko et al., 1974. Rev. Centro Ciências Rurais 4(2), 118; host plant.- Hayes, 1975. Proc. Calif. Acad. Sci. 40(7), 150, fig. 44 (male dorsal); syst.- Chang and Sales, 1977. J. Insect Physiol. 23, 159; biol.- Franclemont and Todd, 1983. Check List Lep. Amer. N. Mexico, 150.- Gonçalves, 1985. Rev. Bras. Ent. 29, 31, figs 1, 2 (digestive tube); morph. - Poole, 1989. Lep. Cat. (n.s.) 118, Noctuidae 2, 844.- Buainain and Silva, 1990. Pesqui. Agropecu. Bras. 25(2), 229; biol., host plant.- Salvadori and Parra, 1990a. Pesqui. Agropecu. Bras. 25(12), 1693; apl., biol.- Salvadori and Parra, 1990b. Pesqui. Agropecu. Bras. 25(12), 1679; apl., biol.- Salvadori and Parra, 1990c. Pesqui. Agropecu. Bras. 25(12), 1701; apl., biol.- Dickel, 1991. Tropical Lep. 2(1), 57, fig. 26 (male); geogr. distr.- Frank and McCoy, 1992. Florida Entomologist 75(1), 12; geogr. distr.- Giannotti et al., 1995. An. Soc. Ent. Brasil 24(3), 460; parasit.- Foerster, 1996. An. Soc. Ent. Brasil 25(1), 27; biol.- Specht and Corseuil, 1996. Biociências 4(2), 146; inv., syst.- Matrangolo et al., 1997. Pesqui. Agropecu. Bras.. 32(8), 773; host plant.- Polaszek and Foerster, 1997. An. Soc. Ent. Brasil 26(1), 180; parasit.- Doetzer and Foerster, 1998. An. Soc. Ent. Brasil 27(2), 255; apl., parasit.- Ferreira, 1998. EMBRAPA-CNPAF, 80, figs 73 (adult), 75 (larva); apl.- Fontana et al., 1998. Appl. Biochem. Biotech. 70-72, 74; apl.- Yamamoto et al., 1998. Acta Biol. Par. 27(1-4), 85; parasit.- Foerster et al., 1999a. An. Soc. Ent. Brasil 28(2), 243-249; apl., parasit.- Foerster et al., 1999b. An. Soc. Ent. Brasil 28(3), 485; apl., parasit.- Specht and Corseuil, 2002a. Pesqui. Agropecu. Bras.. 37(1), 1; inv., host plant.- Specht and Corseuil, 2002b. Rev. Bras. Zool. 19(Supl.1), 286; inv.- Bittencourt et al., 2003. Arq. Inst. Biol. 70(1), 86; inv.- Ohashi and Urdampilleta, 2003. RIA 32(2), 121; ecol., parasit.- Specht et al., 2004. R. Bras. Agrociencia 10(4), 398; inv., host plant.- Specht et al., 2005. Rev. Bras. Ent. 49(1), 136; inv.- Grego et al., 2006. Sci. Agric. 63(4), 321; ecol., apl., host plant.- Torres et al., 2006. CAB Reviews 1(15), 5; apl.- Almeida et al., 2008. Ethol. 114:,449; biol.- Fritz et al., 2008. Oecol. Bras. 12(4), 723; apl., host plant.- Pereira et al., 2008. Braz. Arch. Biol. Technol. 51(2), 259, fig. 1b (parasitismo).- Conceição and Silva, 2011. Campo Digit@l 6(1), 23; apl.- Marchioro and Foerster, 2012a. Agr. For. Entomol. 14, 29; apl., biol., host plant.- Marchioro and Foerster, 2012b. Neotrop. Entomol. 41, 288; apl., biol.- Queiroz et al., 2012. Rev. Bras. Zootec. 41(2), 273; host plant.- Fritz et al., 2013. Inter. Jour. Trop. Ins. Sc. 33(3), 179; apl.- Almeida et al., 2014. Braz. J. Biol. 74(4), 985; inv.- Foerster et al., 2015. BioControl 60, 9; apl., parasit.

Cirphis sequax; Corseuil and Cruz, 1975. Rev. Facul. Agron. UFRGS 1(1), 19; inv.- Specht and Corseuil, 1996. Biociências 4(2), 146; inv., syst.

Mythimna (Pseudaletia) sequax; Foerster et al., 2001. Acta Biol. Par. 30(1-4), 139; apl., parasit.- Foerster and Doetzer, 2002. Neotrop. Ent. 31(3), 405; apl., parasit.- Hacker et al., 2002. Noctuidae Europaeae 4, 168.- Foerster and Doetzer, 2003. Neotrop. Ent. 32(1), 81; apl., parasit.- Yamamoto and Foerster, 2003. Neotrop. Ent. 32(3), 481; parasit.- Barbut and Lalanne-Cassou, 2010. L’Ent. 66(3), 117, figs 3, 4 (male, female), 6, 8 (male and female gen.); syst.- Soria and DeGrande, 2011. Rev. Bras. Milho Sorgo 10(2), 96; inv., host plant.

Mythimna sequax; Lafontaine and Schmidt, 2010. Zookeys 40, 91; syst.- Bortolotto et al., 2015. Cienc. Rural 45(12), 2121; host plant.

Systematic history

Described in Pseudaletia by Franclemont (1951), as it has been most often cited until now.

Type material

Pseudaletia sequax Franclemont, 1951. Species described based on the male holotype from Jalapa, Veracruz, Mexico, deposited at the USNM, with the following labels: /Jalapa, Mex[ico], W. Schaus. / HOLOTYPE Pseudaletia sequax J. G. Franclemont/ USNMENT 00973327/.

Diagnosis

Both sexes with forewings beige or orange-beige, mixed with dark scales similar to those of Mythimna (P.) unipuncta, differing from the latter in the genitalia. In males, the distal third of the cucullus with dorsal margin bent toward the dorsum, ending in a unique scythe-shaped distal process, vesica with basal diverticulum bearing one apical cornutus, raely up to four cornuti apart from each other.

Description

MALE. Head: From beige to orange-neige, mixed with dark scales of different hues. Eyes brown, pilose. Antenna filiform, triangular in cross section, covered dorsally with beige scales, ventrally naked, with tiny chemoreceptive trichoid sensilla, more concentrated in the central part of the antennomer, and with longer lateral chemoreceptive trichoid sensilla, almost as long as antennomer width. Thorax: Ground color as head; patagium with dark subapical line. Forewing: length 1.5-2.2cm (n=51). Ground color from beige to orange-beige, mixed with dark scales of different hues; postmedial line as dark spots over the veins and terminal line as dark spots between veins, other transversal lines absent; with an oblique line of dark scales from the apex to M₂; a dark patch with a spot of central white scales next to the bases of M₂ and M₃; dark patch covering cubital veins absent on most specimens orbicular and reniform spots circular, inconspicuous, orange with smooth darker center; subterminal area of the wing with white or light beige scales over the veins; fringe beige with dark medial and terminal lines. Hindwing pale, with marginal brown shading toward the base, fringe pale. Abdomen: Greyish-beige dorsally; ventrally as head, with two black transversal lines subventral. Genitalia: Tegumen and uncus fused, the former wider than the base of uncus; ventral arms slim in posterior view, covered with setae near the articulation with the dorsal arm of saccus. Saccus bent toward the ventral arms of the tegumen, irregular distally, twisted toward its inner surface. Uncus elongated, flat, and bent ventrally, widened on medial area, apical half abruptally narrowed to 1/5-1/6 of its width, covered with setae dorsally. Valva subtriangular; sacculus projected ventrally, forming an oval lobe; cucullus elongated, apical third with the dorsal margin bent dorsally and ending in a unique scythe-shaped distal process, corona present, with multiple rows of spines; clasper long and oval; ampulla distinct as a digitiform posterior projection; digitus slim, one third or less as long as clasper. Fultura inferior (=juxta) subtriangular, approximately one third the length of the ventral arms of the tegumen and aproximatelly seven times as long as its smallest width, distal portion bifid. Aedeagus pipe-shaped; vesica tubular and elongate, of similar width throughout its length, with a continuous band of cornuti from the opposite side of the diverticulum to subapical area, basal diverticulum elongate, with a single cornutus, rarely with two, three, or four cornuti spaced among them.

FEMALE. Similar to male, but antenna with tiny chemoreceptive trichoid sensilla only, longer lateral chemoreceptive trichoid absent. Forewing length 1.6-2.2cm (n=40). Abdomen: Genitalia almost entirely sclerotized. Eighth tergite straight and narrow. Papilla analis subtriangular covered with setae. Lamella antevaginalis subrectangular. Ductus bursae sclerotized, with a narrow bifurcation at its middle portion, conducting to appendix bursae; corpus bursae membranous and round; appendix bursae with approximately twice the length of ductus bursae, but with the same width; sclerotized basally and with membranous apex. Ductus seminalis originating on appendix bursae apex.

Geographic distribution

Mythimna (P.) sequax is common in the American tropics and subtropics, occurring from Southern U.S.A. to Northern Argentina (Franclemont, 1951) and Uruguay (Biezanko and Ruffinelli, 1971). In Brazil, there are records for most states: Roraima: Alto Alegre; Acre: Rio Branco; Mato Grosso: Sinop; Mato Grosso do Sul; Chapadão do Sul and Miranda; Tocantins: Porto Nacional; Distrito Federal: Planaltina; Pernambuco: Petrolina; Bahia: Correntina and São Desidério; Minas Gerais: Santana do Riacho and Uberaba; Espírito Santo: Alegre and Domingos Martins; Rio de Janeiro: Cachoeiras de Macacu, Itatiaia, Petrópolis and Rio de Janeiro; São Paulo: Cajati, Campos do Jordão, Jundiaí and São Bernardo do Campo; Paraná: Cornélio Procópio, Curitiba, Londrina, Morretes, Ponta Grossa and Quatro Barras; Rio Grande do Sul: Bagé, Passo Fundo, Salvador do Sul and Vacaria.

Host plants

Native and cultivated grasses (Poaceae) such as rice, oat, barley, wheat, corn, canary grass, annual ryegrass, rye, sugar cane, sword grass and other grasses (Biezanko et al., 1974; Buainain and Silva, 1990; Matrangolo et al., 1997; Specht and Corseuil, 2002a; Specht et al., 2004; Fritz et al., 2008; Bortolotto et al., 2015).

Discussion

The coloration of Mythimna (P.) sequax is variable; the dorsal forewing may be beige to orange-beige, variably mixed with dark scales. This species has been frequently confused with Mythimna (P.) unipuncta and Mythimna (P.) celiae sp. nov., but it can be distinguished by the genitalia structures, especially the male genitalia. The vesica has also a basal elongate diverticulum, but in the former it bears a single cornutus or rarely two, three or four, spaced among them, while in the other two it has more than four cornutus.

Material examined

BRAZIL - Roraima: Alto Alegre - 2°56'18.1”S, 61°00'18.0”W, 87m, 1 male, 14-VII-2015, E. G. F. de Morais leg., DZ 39.946 (DZUP). - Acre: Rio Branco - 10°01'57.13”S, 67°37'36.87”W, 207m, 2 males, 16-VII-2015, M. Fazolin leg., DZ 37.899 (DZUP, CPAC), 10°01'58.09”S, 67°42'12.74”W, 183m, 1 female, 5-V-2016, M. Fazolin leg., DZ 39.963 (DZUP). - Tocantins: Porto Nacional - 10°30'36.678”S, 48°18'50.886”W, 212m, 1 female, 8-X-2015, B. A. Evangelista & D. B. Fragoso leg. (CPAC). - Pernambuco: Petrolina - 9°03'54.2”S, 40°10'23.6”W, 365,9m, 1 female, 9-XI-2015, E. A. dos Santos leg. (CPAC), 2 males, 10-XI-2015, E. A. dos Santos leg. (CPAC), 1 female, 13-XI-2015, E. A. dos Santos leg., DZ 39.958 (DZUP), 09°08'14.4”S, 40°18'07.4”W, 365,5m, 1 male, 12-IX-2015, J. B. G. Santos Filho leg., DZ 39.957, 1 male, 13-IX-2015, J. B. G. Santos Filho leg., DZ 39.954 (DZUP). - Mato Grosso: Sinop - 11°52'1.50”S, 55°36'2.19”W, 362m, 1 female, 1 male, 10-VI-2015, A. L. Filipiake, S. M. M. Rodrigues & R. M. Pitta leg. (CPAC), 1 male, 11-VI-2015, A. L. Filipiake, S. M. M. Rodrigues & R. M. Pitta leg., DZ 39.951, 1 female, 13-VI-2015, A. L. Filipiake, S. M. M. Rodrigues & R. M. Pitta leg., DZ 39.952 (DZUP). - Distrito Federal: Planaltina - 15°36'07.10”S, 47°42'46,67”W, 1007m, 1 male, 02-XI-2013, Specht leg. (CPAC), 1 male, 4-XII-2013, A. Specht leg., DZ 39.938 (DZUP), 1 male, 2-II-2014, A. Specht leg. (CPAC), 1 male, 27-IV-2014, A. Specht leg., DZ 39.934, 1 male, 28-IV-2014, A. Specht leg., DZ 39.947 (DZUP), 1 male, 4-V-2014, A. Specht leg. (CPAC), 1 male, 17-VIII-2014, A. Specht leg., DZ 39.940 (DZUP), 15°36'24.52”S, 47°44'42.45”W, 1169m, 1 male, 10-VI-2015, A. Specht leg. (CPAC), 1 female, 10-III-2016, A. Specht leg., DZ 39.936 (DZUP). - Bahia: Correntina - 13°45’24”S, 46°10’01”W, 970m, 1 male, 15-VI-2015, S. V. Paula-Moraes leg., DZ 39.939, 1 male, 15-IX-2015, S. V. Paula-Moraes leg., DZ 39.942 (DZUP); São Desidério - 13°18’60”S, 45°58’ 24”W, 879m, 1 male, 17-III-2015, C. Ceolin leg. (CPAC). - Mato Grosso do Sul: Chapadão do Sul - 18°46'49.6”S, 52°31'1.55”W, 806m, 1 female, 12-VI-2015, T. Taira leg., DZ 39.941 (DZUP), 1 female, 10-IX-2015, T. Taira leg. (CPAC), 1 female, 5-IV-2016, T. Taira leg., DZ 39.962 (DZUP), 1 male, 10-VII-2015, T. Taira leg. (CPAC), 18°46'30.09”S, 52°31'04.98”W, 804m, 1 male, 11-VII-2015, T. Taira leg. (CPAC); Miranda - 20°07'24,29”S, 56°37'15,66”W, 122m, 1 male, 12-VII-2015, H. V. B. Concone leg., DZ 39.923 (DZUP), 20°06'20,20”S, 56°36'52,62”W, 121m, 1 female, 4-II-2016, H. V. B. Concone leg. (CPAC), 1 male, 31-V-2016, H. V. B. Concone leg., DZ 39.945 (DZUP), 2 males, 1-VI-2016, H. V. B. Concone leg., DZ 39.960, DZ 46.908 (DZUP), 1 male, 2-VI-2016, H. V. B. Concone leg., DZ 39.959 (DZUP). - Minas Gerais: Santana do Riacho - Reserva Vellozia, 19°17'00”S, 43°35'34”W, 1100m, 1 male, 9-III-2017, M. Savaris leg., DZ 39.928 (DZUP); Uberaba - 19°39'45.07”S, 47°57'39.16”W, 784m, 1 male, 9-X-2015, B. M. Moreira leg., DZ 39.950 (DZUP), 1 male, 7-I-2016, B. M. Moreira leg. (CPAC), 19°39'14.36”S, 47°58'11.45”W, 819m, 1 female, 7-VII-2015, B. M. Moreira leg. (CPAC), 1 female, 16-XII-2015, B. M. Moreira leg., DZ 39.948 (DZUP). - Espírito Santo: Alegre - 20°45'11,63”S, 41°29'23,28”W, 121m, 1 female, 1 male, 17-VI-2015, L. Mardgan, D. Pratissoli, J. R. Carvalho leg. (CPAC), DZ 39.961, 1 female, 9-X-2015, L. Mardgan, D. Pratissoli, J. R. Carvalho leg., DZ 39.930 (DZUP); Domingos Martins - 20°22'17.3”S, 41°03'47.7”W, 950m, 1 male, 8-III-2016, J. S. Zanuncio Junior leg., DZ 39.932, 1 female, 4-IV-2016, J. S. Zanuncio Junior leg., DZ 39.931 (DZUP). - Rio de Janeiro: Cachoeiras de Macacu - 1 female, 16-IX-2012, N. Tangerini leg., DZ 39.964 (DZUP); Itatiaia - 1 male, 12-13-IX-1950, Travassos, Albuquerque & Silva leg. (IOC), Parque Nacional do Itatiaia, 840m, 1 male, 8-XI-2011, Expedição Lab. Lep. MZUSP leg., 2410m, 2 females, 1 male, 16-17-III-2013, Silva, Pinheiro & Muñoz leg. (MZUSP); Petrópolis - Independência, 1 female, 1930, Ferreira D’Almeida leg., ex. coll. Gagarin DZ 37.876, 1 female, 19-II-1948, Gagarin leg., ex. coll. Gagarin DZ 37.878 (DZUP); Rio de Janeiro - Tijuca, 1 male, 10-X-1916, Ferreira D’Almeida leg., DZ 37.875 (DZUP). - São Paulo: Cajati - Parque Estadual Rio Turvo, Mirante do Aleixo, 24°43'48.081”S, 48°06'29.754”O, 1018m, 1 female, 1-3-XII-2013, Quintero & Silva leg. (MZUSP); Campos do Jordão - Parque Estadual Campos do Jordão, Mirante do Pau Arcado, 22°44'34.322”S, 45°35'46.88”O, 1815m, 1 male, 1 female, 18-19-V-2015, Quintero, Silva & Moraes leg. (MZUSP); Jundiaí - Base Ecológica Serra do Japi, 23°11'8.547”S, 46°53'52.1”W, 3 females, 19-21-X-2012, Expedição Lab. Lep. MZUSP leg. (MZUSP); São Bernardo do Campo – Alto da Serra, 1 female, 1 male, 1927 (MZUSP). - Paraná: Cornélio Procópio - Parque Estadual Mata São Francisco, 23°08'59”S, 50°34'19”W, 656m, 1 male, 3-5-V-2016, Santos, Silva & Queiroz leg., DZ 37.884 (DZUP); Curitiba - 900m, 1 male, 1 female, IX-2004, H. A. Vargas leg., DZ 37.887, DZ 37.888 (DZUP); Londrina - 23°11'44.1”S, 51°10'33.5”W, 594m, 1 female, 10-VII-2015, D. R. Sosa-Gomez leg. (CPAC), 1 female, 10-VIII-2015, D. R. Sosa-Gomez leg. (CPAC), 1 female, 15-IX-2015, Sosa-Gomez leg., DZ 39.956 (DZUP), 23°11'22.9”S, 51°10'18.7”W, 545m, 1 male, 18-IX-2015, D. R. Sosa-Gomez leg., 1 male, 10-XI-2015, Sosa-Gomez leg. (CPAC), 1 male, 6-I-2016, Sosa-Gomez leg., DZ 39.921 (DZUP); Morretes - Parque Estadual Pico Marumbi, Pedra Lascada, 25°26'18,4”S, 48°55'09.6”W, 510m, 2 males, 1 female, 29-IX-3-X-2013, Silva leg. (MZUSP), Estrada da Graciosa, 25°20'2.68”S, 48°53'54.44”W, 700m, 2 males, 14-VII-2017, Dolibaina, C. Mielke, Siewert & St Laurent leg., DZ 39.919, DZ 39.917 (DZUP); Ponta Grossa - 1 female, XI-1955, DZ 37.889, 1 male, XII-1958, ex. coll. F. Justus DZ 37.885 (DZUP); Quatro Barras - Banhado, 1 male, 5-VI-1970, Becker & Laroca leg., DZ 37.883, 1 female, 28-XII-1970, Becker & Laroca leg., DZ 37.882 (DZUP); Tibagi - Parque Estadual do Guartelá, 24°33'44”S, 50°15'33”W, 950-1000m, 5 males, 4 females, 25-27-VII-2017, Dantas, Dias, Dolibaina & Queiroz-S[antos] leg., DZ 39.922, DZ 39.920, DZ 39.933, DZ 39.943, DZ 39.944, DZ 39.916, DZ 39.924, DZ 39.927, DZ 39.937 (DZUP). - Rio Grande do Sul: Bagé - 31°21'4.94”S, 54°1'12.51”W, 232m, 1 female, 8-XII-2015, R. N. Systi, J. U. P. Corrêa, M. A. P. da Silva & Harry Ebert leg. (CPAC); Passo Fundo - 28°13'50.67”S, 52°24'17.04”W, 671m, 1 female, 6-X-2015, P. R. V. S. Pereira leg., DZ 39.955, 1 male, 6-XI-2015, P. R. V. S. Pereira leg., DZ 39.926, 1 female, 12-XII-2015, P. R. V. S. Pereira leg., DZ 39.929, 28°13'35.88”S, 52°24'13.05”W, 682m, 1 male, 6-I-2015, P. R. V. S. Pereira leg., DZ 39.949 (DZUP), 1 male, 11-X-2015, P. R. V. S. Pereira leg. (CPAC), 1 male, 1 female, 9-XII-2015, P. R. V. S. Pereira leg. (CPAC), DZ 39.720, 1 male, 14-II-2016, P. R. V. S. Pereira leg., DZ 39.925 (DZUP), 1 female, 11-II-2016, P. R. V. S. Pereira leg., 1 female, 11-III-2016, P. R. V. S. Pereira leg., 1 male, 12-III-2016, P. R. V. S. Pereira leg. (CPAC); Salvador do Sul - 1 female, 8-XI-1993, A. Specht leg., 2546 (MCTP), 1 female, 4-X-1995, A. Specht leg., 4497 (MCTP); Vacaria - 1 male, 12-II-2009, A. C. Formentini leg.; 1 male, 25-II-2009, A. C. Formentini leg. (CEUCS).

Mythimna (Pseudaletia) celiae sp. nov. (Figs. 1, 12)

Figure 12
Mythimna (P.) celiae sp. nov. a-b, male (DZ 39.913) in dorsal and ventral views; c-d, female (DZ 39.911) in dorsal and ventral views. Specimen deposited at the DZUP. Scale bar = 1cm.

urn:lsid:zoobank.org:pub:D5D278A9-5ADB-476F-BB20-08FA292D1434

Diagnosis

Both sexes with forewing ground color cream, usually mixed with dark scales, on most specimens without dark patch covering cubital vein. In males, cucullus with dorsal margin straight, ending in at least two distal processes, ampulla distinct, as a small ear-shaped dorsal projection, digitus slim, between a third of or as long as the clasper; vesica basal diverticulum elongate, with over four cornuti close to each other. Genitalia similar to Mythimna (P.) unipuncta, differing slightly in external morphology, DNA Barcode (COI) and specific localities.

Description

MALE. Head: Cream, sometimes mixed with dark scales of different hues. Eyes brown, pilose. Antenna filiform, triangular in cross section, covered dorsally with beige scales, ventrally naked, with tiny chemoreceptive trichoid sensilla, more concentrated in the central part of the antennomer and with longer lateral chemoreceptive trichoid sensilla, half as long as antennomer width. Thorax: Ground color as head; patagium with dark subapical line. Forewing: length - 1.5-1.9cm (n=20). Ground color cream, varibale mixed with dark scales of different hues; postmedial line as dark spots over the veins and terminal line as dark spots between veins, other transversal lines absent; with an oblique line of dark scales from the apex to M₂, on some specimens with an anterior area lighter than ground color; a dark patch with a spot of central white scales next to the bases of M₂ and M₃, undifferentiable in many specimens; dark patch covering cubital veins absent on most specimens; orbicular and reniform spots circular, inconspicuous, orange with smooth darker center, undifferentiable in some specimens; subterminal area of the wing with white or light beige scales over the veins; fringe beige with dark medial and terminal lines. Hindwing pale, with marginal brown shading toward the base, fringe pale. Abdomen: Greyish-cream dorsally; ventral view as head, with two faint black transversal lines subventral. Genitalia: Tegumen and uncus fused, the first wider than the base of uncus; ventral arms of tegumen slim, covered with setae posteriorly, close to the articulation with the dorsal arms of saccus. Saccus subquadrangular and bent toward the ventral arms of the tegumen, irregular distally, twisted toward its inner surface. Uncus elongated, flat and bent ventrally, widened on medial area, apical half abruptally narrowed to 1/5-1/6 of its width, covered with setae dorsally. Valva subtriangular; sacculus projected ventrally forming an ovallobe; cucullus long, with a straight dorsal margin and ending in at least two distal processes, distal portion lobate, corona present, with multiple rows of spines; clasper short; ampulla distinct, as a small ear-shaped dorsal projection; digitus slim, 1/3 as long as the clasper, on some speciemens it could be as long as clasper. Fultura inferior (=juxta) subtriangular, approximately one fourth the length of the ventral arms of tegumen and thrice as long as its smallest width, distal portion bifid. Aedeagus pipe-shaped; vesica tubular and elongate, slightly widened apically, with a continuous band of cornuti from posterior the diverticulum to subapical area, basal diverticulum elongate, with over four closely arranged cornuti, the distalmost thinner and longer than the others.

FEMALE. Similar to male, but antenna with tiny chemoreceptive trichoid sensilla only, longer lateral chemoreceptive trichoid absent. Forewing length 1.6-1.9cm (n=14). Abdomen: Genitalia almost entirely sclerotized. Eighth tergite straight and narrow. Papilla analis subtriangular covered with setae. Lamella antevaginalis subrectangular. Ductus bursae sclerotized, with a wide fold at its final portion leading to appendix bursae; corpus bursae membranous and round; appendix bursae with at least twice the length of ductus bursae, but with the same width, sclerotized basally and with membranous apex. Ductus seminalis originating on appendix bursae apex.

Geographic distribution

In Brazil, it is endemic to Rio Grande do Sul State, with records from Bagé, Passo Fundo, and Vacaria municipalities. It probably occurs in Uruguay and Northern Argentina as well, in view of the records of Mythimna (P.) unipuncta to these localities that are probably misidentifications.

Host plants

Unknown. Presumably grasses (Poaceae), as the other species in this study.

Etymology

Homage to Célia Pedrotti Madruga, the first author’s beloved grandmother, whose life was spent in the same meadows where this moth occurs.

Discussion

Mythimna (P.) celiae sp. nov. has the forewings cream colored, variable mixed with dark scales of different hues, but considerably lighter than the other species and the cubital vein not covered by a dark patch. However, this is a variable character, rendering it not the most reliable character for the species identification. Regarding the genitalia, the species can be easily distinguished from Mythimna (P.) adultera and from Mythimna (P.) sequax, with no need of dissection; the former presents the final portion of the dorsal margin of the cucullus protuberant, while in the latter it is bent dorsally, while Mythimna (P.) celiae sp. nov. has the dorsal margin of the cucullus straight, similarly to Mythimna (P.) unipuncta. It can be distinguished from the latter in details, such as the presence of more than one distal process at the tip of the cucullus. During the examination of the material from collections, we found many specimens identified as Mythimna (P.) unipuncta, that are, in fact, Mythimna (P.) celiae sp. nov.

Comparisons were made between specimens of Mythimna (P.) celiae sp. nov. and the descriptions of Mythimna (P.) punctulata (Blanchard, 1852) that occurs in Chile (Blanchard, 1852; Butler, 1882; Franclemont, 1951; Angulo and Weigert, 1977; Parra et al., 1986; Poole, 1989; Artigas, 1994; Olivares et al., 2009) and, possibly, Argentina (Artigas, 1994; Pastrana, 2004), Brazil and Peru (Artigas, 1994). However, the comparative study both of the external morphology and the genitalia (Franclemont, 1951; Angulo and Weigert, 1977), indicated that none of the examined specimens are Mythimna (P.) punctulata. This reinforces the need for more comprehensive studies to elucidate whether the geographic distribution of Mythimna (P.) punctulata is restricted to Chile, or also covers eastern South America, as described in Artigas (1994).

Considering that Mythimna (P.) adultera, Mythimna (P.) sequax and Mythimna (P.) celiae sp. n. are sympatric in Rio Grande do Sul, Brazil, it stands to reason that the identification of the first two may be mistaken (ex. Biezanko et al., 1949; Biezanko and Bertholdi, 1951; Bertels, 1956; Corseuil, 1958; Costa, 1958; Bertels and Baucke, 1966). Likewise, the records for the neighboring countries Uruguay and Argentina of Mythimna (P.) unipuncta and/or Mythimna (P.) punctulata (Artigas, 1994; Pastrana, 2004) may be misidentifications.

In this work, the estimates of evolutionary divergence between the species have shown higher interspecific than intraspecific divergence, and both the tree build using neighbor-joining and the one using maximum likelihood distinguish Mythimna (P.) celiae sp. nov. from Mythimna (P.) unipuncta. Thus, despite the low sample number, molecular, morphological, and geographic distribution data contribute integratively to the diagnosis of Mythimna (P.) celiae sp. nov.

Type material

Male holotype deposited at the DZUP, with the following labels: /HOLOTYPUS/ Brasil, Rio Grande do Sul, Bagé, 31°21'4.94”S, 54°1'12.51”W, 232m, 06-XI-2015, R. N. Systi, J. U. P. Corrêa, M. A. P. da Silva & Harry Ebert leg./ DZ 39.913 / Holotypus Mythimna (Pseudaletia) celiae Madruga, Specht, San Blas, Mielke & Casagrande, 2022/

Female allotype deposited at DZUP, with the following labels: /ALLOTYPUS/ Brasil, Rio Grande do Sul, Bagé, 31°21'4.94”S, 54°1'12.51”W, 232m, 06-XI-2015, R. N. Systi, J. U. P. Corrêa, M. A. P. da Silva & Harry Ebert leg./ DZ 39.911 / Allotypus Mythimna (Pseudaletia) celiae Madruga, Specht, San Blas, Mielke & Casagrande, 2022/

Paratypes: BRAZIL - Rio Grande do Sul: Bagé - 31°18'57.08”S, 53°59'52.91”W, 242m, 1 female, 4-I-2016, R. N. Systi, J. U. P. Corrêa, M. A. P. da Silva & Harry Ebert leg., DZ 39.918, 1 male, 11-III-2016, R. N. Systi, J. U. P. Corrêa, M. A. P. da Silva & Harry Ebert leg., DZ 39.968 (DZUP), 31°21'4.94”S, 54°1'12.51”W, 232m, 1 female, 11-VI-2015, R. N. Systi, J. U. P. Corrêa, M. A. P. da Silva & Harry Ebert leg. (CPAC), 1 male, 11-VIII-2015, R. N. Systi, J. U. P. Corrêa, M. A. P. da Silva & Harry Ebert leg., DZ 39.966, 2 females, 1 male, 12-VIII-2015, R. N. Systi, J. U. P. Corrêa, M. A. P. da Silva & Harry Ebert leg., DZ 39.985 (DZUP, CPAC), 1 male, 13-VIII-2015, R. N. Systi, J. U. P. Corrêa, M. A. P. da Silva & Harry Ebert leg., DZ 37.891 (DZUP), 1 male, 14-VIII-2015, R. N. Systi, J. U. P. Corrêa, M. A. P. da Silva & Harry Ebert leg. (CPAC), 2 females, 7-X-2016, R. N. Systi, J. U. P. Corrêa, M. A. P. da Silva & Harry Ebert leg., DZ 39.984 (DZUP, CPAC), 1 female, 6-XI-2015, R. N. Systi, J. U. P. Corrêa, M. A. P. da Silva & Harry Ebert leg. (CPAC), 1 female, 6-XII-2015, R. N. Systi, J. U. P. Corrêa, M. A. P. da Silva & Harry Ebert leg. (CPAC); Passo Fundo - 28°13'35.88”S, 52°24'13.05”W, 682m, 1 female, 10-VIII-2015, P. R. V. S. Pereira leg., DZ 39.967, 1 male, 8-IX-2015, P. R. V. S. Pereira leg., DZ 37.890, 1 male, 8-X-2015, P. R. V. S. Pereira leg., DZ 39.988 (DZUP), 3 males, 1 female, 9-X-2015, P. R. V. S. Pereira leg. (CPAC), DZ 39.914, 2 females, 2 males, 11-X-2015, P. R. V. S. Pereira leg., DZ 39.905, DZ 39.915, DZ 39.904, DZ 39.907, 1 female, 2 males, 7-XI-2015, P. R. V. S. Pereira leg., DZ 39.987 (DZUP, CPAC), 1 male, 1 female, 9-XII-2015, P. R. V. S. Pereira leg., DZ 39.980 (DZUP, CPAC), 2 males, 1 female, 11-XII-2015, P. R. V. S. Pereira leg. (CPAC), 1 female, 5-I-2016, P. R. V. S. Pereira leg., DZ 39.982 (DZUP), 28°13'50.67”S, 52°24'17.04”W, 671m, 1 female, 9-X-2015, P. R. V. S. Pereira leg.(CPAC), 1 female, 10-X-2015, P. R. V. S. Pereira leg., DZ 39.986 (DZUP), 3 males, 5-XI-2015, P. R. V. S. Pereira leg. (CPAC), DZ 39.908, DZ 39.909 (DZUP), 1 female, 1 male, 9-XI-2015, P. R. V. S. Pereira leg. (CPAC), DZ 39.910 (DZUP), 2 females, 12-XII-2015, P. R. V. S. Pereira leg., DZ 39.912 (DZUP, CPAC), 1 female, 13-II-2016, P. R. V. S. Pereira leg., DZ 39.906, 1 male, 2 females, 11-II-2016, P. R. V. S. Pereira leg., DZ 39.983, DZ 39.981 (DZUP, CPAC), 1 male, 9-III-2016, P. R. V. S. Pereira leg. (CPAC); Vacaria - 1 male, 8-II-2009 (CEUCS), 1 female, 12-II-2009 (CEUCS), 1 female, 19-II-2009 (CEUCS), 1 male, 25-II-2009 (CEUCS).

Final considerations

The examination of the specimens available at several collections corroborated the understanding that, in Brazil, over the years, many individuals of Mythimna (Pseudaletia) were incorrectly identified as Mythimna (Pseudaletia) unipuncta, due to the fact that they form a complex of externally very similar species. This reinforces the need for the genitalia examination to achieve a more reliable specific identification. In this sense, even though the study of genitalia characters is a recurrent practice in the taxonomy of the group (Yoshimatsu, 1994, 1995; Yoshimatsu and Hreblay, 1998; Yoshimatsu and Legrain, 2001), it is important that, besides making the practical protocols available, publications with images and specific detailed descriptions are provided, including of immatures and adults of each species (Madruga et al., 2019). In Brazil and neighboring countries (CONESUL), given the greater occurrence and importance of these species, publications comparing all development stages of Mythimna (P.) adultera, Mythimna (P.) celiae, and Mythimna (P.) sequax are essential.

Particularly in Rio Grande do Sul, Mythimna (P.) celiae sp. nov. was the species most confused with Mythimna (P.) unipuncta. This probably happened because Gassen (1984) differentiated Mythimna (P.) adultera from Mythimna (P.) sequax. Hence, when researchers could not associate their specimens to one of these two species, they associated the then undescribed species to Mythimna (P.) unipuncta. Since Mythimna (P.) celiae sp. nov. has been collected in several localities in Rio Grande do Sul, including the biome Campos Sulinos, which is contiguous with Uruguay and Argentina, it is likely that the records of Mythimna (P.) unipuncta to this neighboring countries (e.g. Salas et al., 1998; Pastrana, 2004; Isas et al., 2016), are also misidentification. Similarly, the invalid lectotype of Leucania extranea, designated by Viette (IV-1951), which is a synonym of Mythimna (P.) unipuncta, from Montevideo, Uruguay, probably corresponds to Mythimna (P.) celiae sp. nov.

It is important to highlight that, although there has been structured collecting throughout the whole country and an examination of the available material in all Brazilian collections, Mythimna (P.) unipuncta has a single record in Uiramutã, Monte Roraima (Franclemont, 1951), in the extreme North of the country.

Besides morphology and geographic distribution, the species were separated through molecular analyses, DNA Barcode (COI). According to Ball and Armstrong (2006), molecular information may be highly precise, being a promising tool in the identification of pest insects. This method has already been used to differentiate species of Mythimna from Indonesia (Sutrisno, 2012). Moreover, the genetic structure of Mythimna (P.) unipuncta was studied in the archipelago of Azores, Portugal, in continental Portugal, and in Canada, with little differences being found between the populations (Vieira et al., 2003). The estimates of evolutionary divergence and the construction of evolutionary trees (Figs. 3-4), corroborate with morphological and distributional data, confirming that Mythimna (P.) celiae sp. nov. and Mythimna (P.) unipuncta are distinct species.

Mythimna punctulata (Blanchard, 1852) was mentioned for Brazil by Artigas (1994), but based on external morphology and genitalia, their occurrence was not confirmed in this study.

Our results, as suggested in the revision performed by Franclemont (1951), indicate the need of a Neotropical review, especially because of the historical misidentifications related to Mythimna (P.) unipuncta and because of the possible wider geographic distribution of Mythimna (P.) celiae sp. nov. toward Uruguay and Argentina. This full revision would have a local repercussion related to specific recognition and management of these species, as well as an international impact, for Mythimna (P.) unipuncta, considered present in all South American countries (CABI, 2019) in fact is not, its occurrence as a quarantine pest in South America should be reviewed.

In conclusion, the occurrence of the following species is confirmed to Brazil:

Mythimna (Pseudaletia) unipuncta (Haworth, 1809) - Brazil (Roraima)

Leucania extranea Guenée, 1852

Leucania antica Walker, 1856

Pseudaletia unipuncta quechua Franclemont, 1951

Leucania unipuncta tseki Koutsaftikis, 1974

Pseudaletia roraimae Franclemont, 1951, syn. nov.

Mythimna (Pseudaletia) adultera (Schaus, 1894) - Brazil (Paraná, Santa Catarina and Rio Grande do Sul)

Mythimna (Pseudaletia) sequax Franclemont, 1951 - Brazil

Mythimna (Pseudaletia) celiae Madruga, Specht, San Blas, Mielke & Casagrande sp. nov. – Brazil (Rio Grande do Sul).

Acknowledgments

The authors thanks the curators of the cited collections for loan material. To Paul Goldstein (USNM) and Maria Dimanski (Goulandris Museum) for information on type material of Leucania extranea and Leucania unipuncta tseki respectively. To Alfred Moser (CLAM), Amabílio José Aires de Camargo (CPAC), Andrew D. Warren (MGCL), Eduardo J.E. Silva (MECB), David Grimaldi (AMNH), Gervásio Carvalho (MCTP), Jane Costa and Aline V. Miranda (IOC), Paul Goldstein (USNM), Alberto Zilli (NHMUK), Rodolphe Rougerie (MNHN), Wilson Sampaio de Azevedo Filho (CEUCS), Marcelo Duarte and Renato de Oliveira da Silva (MZUSP), Sonia Lopes and Alexandre Soares (MNRJ) for providing access to specimens in their care. To all person involved with field collections of specimens: Adriano Q. Mesquita, Américo I. Ciociola Júnior, André L. Filipiake, Antônio C.S. Araújo, Balbino A. Evangelista, Brenda M. Moreira, Daniel B. Fragoso, Daniel R. Sosa-Gomez, Denivaldo C. Castro, Dirceu Pratissoli, Elisângela G. Fidelis, Erivaldo A. Santos, Felipe O. Mateus, Harry Ebert, João B.G. Santos Filho, Jorge U. P. Corrêa, José A. Teston, José R. Carvalho, José S. Zanúncio Júnior, Leonardo Mardgan, Maicon Coradini, Marco A.P. Silva, Marcos R.O. Serpa, Murilo Fazolin, Naylor B. Perez, Oriverto Tonon, Paulo R.V.S. Pereira, Rafael M. Pitta, Rinaldo J. Silva Júnior, Rodison N. Sisti, Sandra M.M. Rodrigues, Tiago C.C. Lima, Vander C.M. Claudino, and Wilson Pozenato. We are also grateful to ICMBio and MMA for the Authorizations for Scientific Activities (SISBIO no. 48218-3 and 38547/6).

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Edited by

Associate Editor: Héctor Vargas

Publication Dates

• Publication in this collection
  19 Sept 2022
• Date of issue
  2022

History

• Received
  25 Apr 2022
• Accepted
  17 Aug 2022