First description of the later immature stages of Orthos hyalinus (Bell, 1930) (Hesperiinae: Hesperiini: Carystina)

117 First description of the later immature stages of Orthos hyalinus (Bell, 1930) (Hesperiinae: Hesperiini: Carystina) Ayane Suênia-Bastos1,3*, Suianne Cajé1,4, Adalberto Dantas de Medeiros1,5, Emanoel Pereira Gualberto1,6, Jefferson Duarte-de-Mélo2,7, Joshua Pablo Cavalcanti2,8 and Olaf Hermann Hendrik Mielke1,9 1. Laboratório de Estudos de Lepidoptera Neotropical, Departamento de Zoologia, Universidade Federal do Paraná (UFPR), Curitiba, Paraná, Brazil; 2. Departamento de Zoologia, Universidade Federal do Paraná (UFPR), Curitiba, Paraná, Brazil. *Corresponding author: [email protected]. 3. https://orcid. org/0000-0002-3443-745X; 4. https://orcid.org/0000-0002-4558-4551; 5. https://orcid.org/0000-0002-7675-2407; 6. https://orcid.org/0000-0001-9895-9460; 7. https://orcid.org/0000-0003-0268-279X; 8. https://orcid.o.rg/0000-0002-1285-0745; 9. https://orcid.org/0000-0003-3655-4606 Date of issue online: 19 December 2025 Electronic copies (ISSN 2575-9256) in PDF format at: https://journals.flvc.org/troplep; https://zenodo.org; archived by the Institutional Repository at the University of Florida (IR@UF), http://ufdc.ufl.edu/ufir; DOI: 10.5281/zenodo.17860825 © The author(s). This is an open access article distributed under the Creative Commons license CC BY-NC 4.0 (https://creativecommons.org/ licenses/by-nc/4.0/). Abstract: Immatures of Orthos hyalinus (Bell, 1930), found on Phyllostachys aurea Carrière ex Rivière & C. Rivière, an exotic Poaceae, are described and illustrated herein for the first time. The three last instars, the shelter, and the pupa were recorded and are illustrated, and a female emerged from one of the resulting pupae. In addition, a brief discussion of larval and pupal morphology is provided. Key words: food plants; host plants; Neotropical region; Poaceae; taxonomy. Some Hesperiidae subfamilies feed on dicotyledons, whereas others feed on monocotyledons (Warren et al., 2009), and this pattern is the result of two inferred shifts in feeding behaviour from dicotyledons to monocotyledons during the evolution of the family Hesperiidae (Sahoo et al., 2017). Plants have evolved a range of strategies to deter herbivores, from mechanical defenses in their leaf structures to chemical compounds that serve either to act directly on the herbivore or to attract natural enemies (War et al., 2012). Unlike dicotyledons, most Poaceae (a common host plant of the subfamily Hesperiinae) lack the variety of chemical compounds needed to deter herbivores, relying mainly on mechanical defense (Tscharntke & Greiler, 1995; Massey et al., 2006). These differences between major groups of plants may have promoted the diversification of some Hesperiidae lineages; for example, the diversification of Hesperiinae accelerated soon after the shift from one feeding behavior to another, potentially as a result of the opportunity to exploit unoccupied niches (Sahoo et al., 2017). Currently, Hesperiinae is considered the richest subfamily within Hesperiidae, with most of its representatives found in the Neotropical region (Mielke, 2025). However, aspects of the immature stages relevant to the taxonomy and evolution of Hesperiinae, the most diverse subfamily and one of the most conspicuous groups of skippers, remain poorly studied or completely unknown. One main reason for this imbalance is that adults are much easier to collect than larvae, even though immatures may provide important characters to help build higher classifications (Scoble, 1992; Freitas & Brown, 2004; Lin et al., 2019), or to understand evolutionary patterns (Sierra-Botero et al., 2022; Kawahara et al., 2023). Orthos Evans, 1955 (Hesperiini: Carystina) is one of many examples in Hesperiinae for which there is no information on its immature stages or host plant family. Thus, to begin filling these kinds of gaps, this study aims to describe the final instars and pupa of Orthos hyalinus (Bell, 1930), including notes on behavior. During fieldwork, two immatures were found by the first author in February 2025 around the Centro Politécnico at the Universidade Federal do Paraná (UFPR), Curitiba, Paraná, Brazil (25°26'51.2"S, 49°14'12.5"W, 910 m a.s.l.). These immatures, along with the leaves from the host plant, were placed in plastic containers (250 ml) and maintained in the laboratory to allow daily monitoring of development. Leaf shelters were opened after each molt to document each instar. Photographs were taken using a Leica M205C stereomicroscope equipped with Leica DMC4500 digital camera, operated via Leica Application Suite Version 4.12.0 (Build 86), a Sony digital camera, and a Canon digital camera. Measurements were taken with ImageJ software (Schneider et al., 2012) of the head capsule (the widest portion) and body (from head to the last abdominal segment). The terminology of Scoble (1992) was followed to describe the three last larval instars and pupa and Greeney (2009) for describing the leaf shelter. The host plant exsiccate was identified as Phyllostachys aurea Carrière ex Rivière & C. Rivière, an exotic Poaceae, and has been deposited in the Herbarium of Universidade Federal do Paraná (UPCB) under the number UPCB 108434, with images of the voucher specimen available online at JBRJ (https://jabot.jbrj. gov.br/). One larva died during the antepenultimate instar. The other individual completed its development in 75 days from larva to adult emergence. This specimen was deposited in the collection Padre Jesus Santiago Moure, Zoology Department, at the Universidade Federal do Paraná (DZ 73.625). Here, the TROP. LEPID. RES., 35(2): 117-121, 2025SUÊNIA-BASTOS ET AL.: Immatures of Orthos hyalinus 118 TROP. LEPID. RES., 35(2): 117-121, 2025 SUÊNIA-BASTOS ET AL.: Immatures of Orthos hyalinus Figure 1. Three last instars and shelter behaviour of Orthos hyalinus (Bell, 1930). A-B) Antepenultimate instar, dorsal and lateral views. C-D) Penultimate instar, dorsal and lateral views. E-F) Last instar, dorsal and lateral views. G) General view of shelter built by the antepenultimate instar; H) Red arrow indicating the shelter during the last instar. I-J) Pupa, dorsal and lateral view; a silk belt in the thorax reinforcing to the hostplant. morphology of the last three instars, pupa, and shelter behavior is described. Antepenultimate instar (Fig. 1A-B): Head capsule flattened in lateral view, dark creamish, with a black spot in middle of head capsule, containing small dark creamish dots; one brownish band laterally in antero-posterior region; six stemmata, fifth near antennal base. Body with segments T1 to A1 pale greenish, segments A2 to A7 dark greenish, segments A8 to A10 almost whitish; prothoracic shield pale greenish; abdominal spiracles yellowish; last abdominal segment with long setae in distal portion. Head capsule width (n=1): 1.6 mm. Body length (n=1): 12.80 mm. Development time since collecting: 14 days. Penultimate instar (Fig. 1C-D): Head capsule similar to previous instar, but pale creamish, with black spot reduced to three longitudinal lines, inner larger on epicranial suture; same brownish band laterally. Body similar to previous instar, but paler. Head capsule (n=1): 2.1 mm. Body length (n=1): 14.16 mm. Development time: 15 days. Last instar (Fig. 1E-F): Head capsule completely creamish dorsally, without spots or lines near to epicranial suture; same brownish band laterally. Body completely pale greenish, paler dorsally, almost whitish; abdominal spiracles whitish. Head capsule (n=1): 2.7 mm Body length (n=1): 29.09 mm. Development time: 21 days. Shelter behavior (Fig. 1G-H): During all these stages, the larva built shelter type I - no cut (Greeney, 2009), which involved only attaching two leaves with silk, and it stayed inside and ate the leaf of its shelter. The immature pupated inside this shelter. Pupa (Fig. 1I-J): Head capsule greenish with a single long projection anteriorly, arched dorsally in lateral view (Fig. 1J). Body greenish with abdominal segments paler. Pupa attached to leaf by a belt silk on thorax. Female emerged after 15 days. Body length (n=1): 25.34 mm. Although no male emerged, both sexes are illustrated here (Fig. 2A-D), with a male specimen from DZUP collection. 119 Figure 2. Orthos hyalinus (Bell, 1930), dorsal view and ventral view. A, B) Male from Vila Velha (900 m a.s.l), Ponta Grossa, Paraná, Brazil (DZ 73.646). C, D) Female emerged, from Centro Politécnico (UFPR), Curitiba, Paraná, Brazil (DZ 73.625). The original description of Orthos hyalinus (Bell, 1930) was based on a single male collected in Hansa Humboldt [now Corupá] (Santa Catarina, Brazil), and subsequently the species has been recorded in occurrence lists from southeastern and southern Brazil as well as Argentina: Brazil: Espírito Santo (Brown & Freitas, 2000: 113), Rio de Janeiro (Evans, 1955: 293; Zikán & Zikán, 1968: 59), São Paulo (Evans, 1955: 293; Martins, 1993: 121; Francini et al., 2011: 67), Paraná (C. Mielke, 1994: 765; Dolibaina et al., 2011: 345; O. Mielke et al., 2012: 64; Greve et al., 2023: 13), Santa Catarina (Bell, 1930: 152; Hoffman, 1934: 72; Orlandin et al., 2020: 69), Rio Grande do Sul (Biezanko & O. Mielke, 1973: 92; Giovenardi et al., 2013: 15). Argentina: Misiones (Núñez Bustos, 2008: 80; Núñez-Bustos, 2009: 77; Klimaitis et al., 2018: 276). In this paper, the first illustrations and descriptions of the later immature stages, host plant and shelter behavior are provided for an Orthos species. A brief discussion on the morphology of the immatures is also provided here. In terms of knowledge of related species, Corta lycortas (Godman, 1900), previously classified in Orthos, was recorded feeding on Beaucarnea recurvata (K. Koch & Fintelm.) Lem. (Asparagaceae) in Paso del Abra, Tamaulipas (Mexico) (Kendall & McGuire, 1975). The shelter of C. lycortas seems to be similar to that found in O. hyalinus, based on the description of Kendal & McGuire (1975). Additionally, the head capsule of O. hyalinus immatures is curiously distinct from that of other Hesperiinae species (e.g., Freitas, 2020; Orlandin et al., 2020; Ccahuana et al., 2021; Raby et al., 2024; Medeiros et al. 2025; Vega et al., 2025), being so flattened in lateral view that it gives the impression of a prognathous head, although it is hypognathous. Minno (1994) drew attention to modifications in the head of Hesperiidae pupae. The presence of a pupal cephalic projection seems to be an important character found in O. hyalinus and other Neotropical and Neartic Hesperiinae (see Table 1). When present, its form may vary from long to short, and based on the available data for some Hesperiinae, this projection can be classified into three types (see Table 1): (1) straight (not arched); (2) dorsally arched; (3) ventrally arched. Other studies have noted the presence of cephalic projections of pupae from at least six different Hesperiidae subfamilies, with some of these morphologically distinct. Atkins (1978) described similar structures, referred as ‘pupal caps’, in four Australian Hesperilla Hewitson, 1868 (Trapezitinae), and Toussaint et al. (2022) used the same term for Atkinsia dominula (Plötz, 1884) (Trapezitinae). Chiba et al. (1991) described a single, short cephalic projection in the Asian Coladenia minor Chiba, 1991 (Tagiadinae: Tagiadini). Minno (1994) describes the presence of this structure in Neartic Hesperiidae species, naming it a ‘pupal cap’, following Atkins (1978). Cock (2010) provided illustrations of projections in Pyrrhiades anchises anchises (Gerstaecker, 1871) and species of Coeliades Hübner, 1818, both Afrotropical Coeliadinae. Cock & Congdon (2011) illustrated the multiple cephalic projections in Celaenorrhinus Hübner, [1819] (Tagiadinae: Celaenorrhinini) from the Afrotropics. Orlandin et al. (2020) recorded a short cephalic projection in the Neotropical Eantis (Eantis) thraso (Hübner, [1807]) (Pyrginae: Achlyodini: Achlyodina), and what seems to be a beginning of projection in Polygonus leo (Gmelin, [1790]) (Eudaminae: Eudamini: Telemiadina). Cock & Congdon (2017) illustrated cephalic projections similar to Hesperiinae in Afrotropical species of Metisella Hemming, 1934 (Heteropterinae). Finally, Burns et al. (2023) reported an extremely rough and truncate cephalic projection in the Neotropical Ridens conservationinternationalis Burns & Grishin, 2023 (Eudaminae: Eudamini: Loboclina). The pupal cephalic projection has now been documented as present in at least in seven Hesperiidae subfamilies. TROP. LEPID. RES., 35(2): 117-121, 2025SUÊNIA-BASTOS ET AL.: Immatures of Orthos hyalinus 120 Table 1. Examples of Hesperiinae species with pupal cephalic projection, direction of the projection, and the source of information. Tribe Subtribe Species Direction Reference Pericharini Pericharina Orses cynisca (Swainson 1821) Ventrally arched Cock (2005) Oz ozias ozias (Hewitson, 1878) Ventrally arched Freitas (2018); Orlandin et al. (2020) Perichares aurina Evans, 1955 Ventrally arched Orlandin et al. (2020) Perichares geonomaphaga Burns, 2008 Ventrally arched Burns et al. (2008) Perichares philetes (Gmelin, [1790]) Ventrally arched Minno (1994) Perichares poaceaphaga Burns, 2008 Ventrally arched Burns et al. (2008) Perichares prestoeaphaga Burns, 2008 Ventrally arched Perichares lotus (A. Butler, 1870) Ventrally arched Suênia-Bastos et al. (in prep.) Hesperiini Calpodina Calpodes ethlius (Stoll. 1782) Dorsally arched Minno (1994) / Orlandin et al. (2020) Calpodes longirostris (Sepp, [1840]) Straight Freitas (2020) Carystina Carystus (Synale) ploetzi O. Mielke & Casagrande, 2002 Dorsally arched Cock (2005) Cobalus virbius virbius (Cramer, 1777) Dorsally arched Ebusus ebusus ebusus (Cramer, 1780) Straight Ccahuana et al. (2021) Talides sinois Hübner, [1819] Dorsally arched Cock (2005) Talides sergestus (Cramer, 1775) Straight Panoquina panoquin (Scudder, 1863) Straight Minno (1994) Falgina Falga ombra Evans, 1955 Straight Greeney & Warren (2009b) Hesperiina Xeniades (Xeniades) orchamus (Cramer, 1777) Ventrally arched Sherer et al. (2025) Moncina Artonia artona (Hewitson, 1868) Ventrally arched Medeiros et al. (in prep.) Cobalopsis nero (Herrich-Schäffer, 1869) Straight Orlandin et al. (2020) Corra coryna (Hewitson, 1866) Dorsally arched Greeney & Warren (2009a) Cymaenes laureolus (Schaus, 1913) Straight Orlandin et al. (2020) Cymaenes tripunctus theogenis (Capronnier, 1874) Straight Cock (2012) Dion carmenta carmenta (Hewitson, 1870) Straight Greeney & Warren (2011) Dubia dubia (Bell, 1932) Ventrally arched Sherer et al. (2025) Lerema (Lerema) ancillaris (Butler, 1877) Straight Cock (2013) Troyus phyllides (Röber, 1925) Straight Duerr et al. (2022) Vehilius jabre Medeiros, Souza & Kerpel, 2023 Ventrally arched Souza et al. (2023) ACKNOWLEDGMENTS We are grateful to Marília Locatelli Corrêa for depositing and preparing the voucher of P. aurea at the Herbarium of Universidade Federal do Paraná, and Regina Tomoko Shirasuna for the species identification. We thank Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) for fellowships to ASB (88887.949958/2024-00), SC (88887.931876/2024-00), JDM (88887.931875/2024-00), JPC (88887.132404/2025-00), and ADM (88887.941144/202400). The authors acknowledge the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for the research fellowship to EPG (132047/2024-1). Finally, we thank the reviewers who made suggestions that improved this work, and Keith Willmott for editing the manuscript. LITERATURE CITED Atkins, A. 1978. The Hesperilla malindeva group from northern Australia, including a new species (Lepidoptera: Trapezitinae). Journal of the Australian Entomological Society 17: 205-215. Bell, E. L. 1930. Descriptions of new South American Hesperiidae (Lepidoptera, Rhopalocera). Journal of the New York Entomological Society 38: 149158. Biezanko, C. M., Mielke O. H. H. 1973. Contribuição ao estudo faunístico dos Hesperiidae Americanos. IV Espécies do Rio Grande do Sul, Brasil, com notas taxonômicas e descrições de espécies novas (Lepidoptera). Acta Biológica Paranaense 2(1,2,3,4): 51-102. Brown, K. S., Freitas, A. V. L. 2000. Diversidade de Lepidoptera em Santa Teresa, Espírito Santo. Boletin do Museu de Biologia Prof. Mello Leitão (Nova Série) 11/12: 71-118. Burns, J. M., Janzen, D. H., Hajibabaei, M., Hallwachs, W., Hebert, P. D. N. 2008. DNA barcodes and cryptic species of skipper butterflies in the genus Perichares in Area de Conservación Guanacaste, Costa Rica. Proceedings of the National Academy of Sciences of the USA 105(17): 6350-6355. Burns, J. M., Janzen, D. H., Hallwachs, W., Grishin, N. V. 2023. Going for a Ridens Evans (Lepidoptera: Hesperiidae: Eudaminae): a new species reared in Area de Conservacion Guanacaste, northwestern Costa Rica. Proceedings of the Entomological Society of Washington 124(4): 721734. Ccahuana, R., Tejeira, R., Hurtado, T., Nakahara, S., Rodríguez-Melgarejo, M., Gott, R. J., See, J., Gallice, G. 2021. Immature stages of Ebusus ebusus ebusus (Cramer, 1780) in the Peruvian Amazon (Lepidoptera: Hesperiidae: Hesperiinae). Tropical Lepidoptera Research 31(2): 90-95. Chiba, H., Nakanishi, A., Fukuda, H., Yata, O. 1991. A new species of Coladenia from Luzon, Philippines, with description of immatures (Lepidoptera: Hesperiidae). Tropical Lepidoptera Research 2(1): 59-64. Cock, M. J. W. 2005. The skipper butterflies (Hesperiidae) of Trinidad. Part 13, Hesperiinae, Genera Group K. Living World, Journal of the Trinidad and Tobago Field Naturalists’ Club 2005: 23-47. Cock, M. J. 2010. Observations on the biology of Afro-tropical Hesperiidae (Lepidoptera) principally from Kenya. Part 1. Introduction and Coeliadinae. Zootaxa 2547(1): 1-63. Cock, M. J. 2013. The skipper butterflies (Hesperiidae) of Trinidad. Part 19. Hesperiinae, Moncini: the remaining genera with pale spots: Cymaenes, Cobalopsis, Arita, Lerema, Morys and Tigasis. Living World, Journal of the Trinidad and Tobago Field Naturalists’ club 2013: 20-40. Cock, M. J., Congdon, C. E. 2011. Observations on the biology of Afrotropical Hesperiidae (Lepidoptera) principally from Kenya. Part 3. Pyrginae: Celaenorrhinini. Zootaxa 3033(1): 1-67. Cock, M. J., Congdon, C. E. 2017. Observations on the biology of Afrotropical Hesperiidae (Lepidoptera) with particular reference to Kenya. Part 11. Heteropterinae. Zootaxa 4226 (4): 487-508. Dolibaina, D. R., Mielke, O. H. H., Casagrande, M. M. 2011. Borboletas (Papilionoidea e Hesperioidea) de Guarapuava e arredores, Paraná, Brasil: um inventário com base em 63 anos de registros. Biota Neotropica 11(1): 341-354. Duerr, N., Corahua-Espinoza, T., Baine, Q., Tejeira, R., Ccahuana, R., Espinoza, M. M. C., Perlett, E., Cervantes-Martínez, J. N., Santillana, A. L., See, J., Soto-Quispe, Y. S., Wood, H., Arteaga, Z. E., Gallice, G. 2022. Immature stages, new host plant records and shelter structures of Troyus phyllides (Röber, 1925) and Thoon ponka Evans, 1955 in the Peruvian Amazon (Lepidoptera: Hesperiidae: Hesperiinae: Hesperiini). TROP. LEPID. RES., 35(2): 117-121, 2025 SUÊNIA-BASTOS ET AL.: Immatures of Orthos hyalinus 121 Zootaxa 5200 (4): 372-390. Evans, W. H. 1955. A Catalogue of the American Hesperiidae Indicating the Classification and Nomenclature Adopted in the British Museum (Natural History). Part IV. Hesperiinae and Megathyminae. London, British Museum. v + 499 pp., pls. 54-88. Francini, R. B., Duarte, M., Mielke, O. H. H., Caldas, A., Freitas, A. V. L. 2011. Butterflies (Lepidoptera, Papilionoidea and Hesperioidea) of the “Baixada Santista” region, coastal São Paulo, southeastern Brazil. Revista Brasileira de Entomologia 55(1): 55-68. Freitas, A. V. 2018. Immature stages of the Neotropical skipper Lychnuchoides ozias ozias (Hewitson, 1878) (Lepidoptera: Hesperiidae). Tropical Lepidoptera Research 28(1): 25-28. Freitas, A. V. L. 2020. Immature stages of the Neotropical skipper Saliana longirostris (Sepp, [1840]) (Lepidoptera: Hesperiidae). Tropical Lepidoptera Research 30(2): 81-85. Freitas, A. V. L., Brown Jr., K. S. 2004. Phylogeny of the Nymphalidae (Lepidoptera). Systematic Biology 53(3): 363-383. Giovenardi, R., Di Mare, R. A., Mielke, O. H. H., Casagrande, M. M., Carneiro, E. 2013. Mariposas de Rio Grande do Sul, Brasil (Lepidoptera: Papilionoidea, Hesperioidea). Revista Colombiana de Entomología 39(2): 267-275. Greeney, H. F. 2009. A revised classification scheme for larval hesperiid shelter, with comments on shelter diversity in the Pyrginae. Journal of Research on the Lepidoptera 41: 53-59. Greeney, H. F., Warren A. D. 2009a. The life history and shelter building behavior of Vettius coryna coryna Hewitson, 1866 in eastern Ecuador (Lepidoptera: Hesperiidae: Hesperiinae). Journal of Insect Science 9(33): 1-10. Greeney, H. F., Warren, A. D. 2009b. The immature stages and shelter building behavior of Falga jeconia ombra in eastern Ecuador (Lepidoptera: Hesperiidae: Hesperiinae). Journal of Insect Science 9(33): 1-10. Greeney, H. F., Warren, A. D. 2011. The natural history, immature stages, and shelter building behavior of Dion carmenta (Lepidoptera: Hesperiidae: Hesperiinae) in eastern Ecuador. Annals of the Entomological Society of America 104(6): 1128-1134. Greve, R. R., Carneiro, E., Mielke, O. H. H., Robbins, R. K., Callaghan, C. J., Freitas, A. V. L. 2023. Butterflies (Lepidoptera: Papilionoidea) of Iguaçu National Park and surrounding areas in southern Brazil: a longterm survey, with six new records for Brazilian fauna. Biota Neotropica 23(3): e20231487. Hoffmann, F. 1934. Beiträge zur Lepidopterenfauna von St. Catharina (Südbrasilien). Entomologische Rundschau 51: 71-74. Kawahara, A. Y., Storer, C., et al. 2023. A global phylogeny of butterflies reveals their evolutionary history, ancestral host and biogeographic origins. Nature Ecology and Evolution 7: 903-913. Kendall, R. O., McGuire, W. W. 1975. Larval foodplants for twenty-one species of skippers (Lepidoptera: Hesperiidae) from Mexico. Bulletin of the Allyn Museum 27: 1-7. Klimaitis, J. F., Núñez Bustos, E. O., Klimatis, C. L., Güller, R. M. 2018. Mariposas Argentina: Guía de Identificación. Buenos Aires, Vásquez Mazzini. 327 pp. Lin, R.-J., Braby, M. F., Hsu, Y.-F. 2019. The immature stages, biology, and phylogenetic relationships of Rotunda rotundapex (Lepidoptera: Bombycidae). Journal of Insect Science 19(2): 1-11. Martins, R. P. 1993. The biology of Editha magnifica (Perty, 834) (Hymenoptera Sphecidae). Tropical Zoology 6: 109-123. Massey, F. P., Ennos, A. R., Hartley, S. E. 2006. Silica in grasses as a defence against insect herbivores: contrasting effects on folivores and a phloem feeder. Journal of Animal Ecology 75(2): 595-603. Medeiros, A. M., de Medeiros, A. D., Kerpel, S. M. 2025. Immature stages of Callimormus saturnus (Herrich-Schäffer, 1869) (Lepidoptera: Hesperiidae, Hesperiinae, Hesperiini, Moncina): biology, morphology and behaviour. Historia Naturalis Bulgarica 47(6): 145-156. Mielke, C. G. C. 1994. Papilionoidea e Hesperioidea (Lepidoptera) de Curitiba e seus arredores, Paraná, Brasil, com notas taxonômicas sobre Hesperiidae. Revista Brasileira de Zoologia 11(4): 759-776. Mielke, O. H. H., Carneiro, E., Casagrande, M. M. 2012. Hesperiidae (Lepidoptera, Hesperioidea) from Ponta Grossa, Paraná, Brazil: 70 years of records with special reference to faunal composition of Vila Velha State Park. Revista Brasileira de Entomologia 56(1): 59-66. Mielke, O. H. H. 2025. Checklist of the American Hesperiidae. Version August 5, 2025. [26-XI-2025] https://sites.google.com/site/lablepneo/ publica%C3%A7%C3%B5es/livros? Minno, M. C. 1994. Immature stages of the skipper butterflies (Lepidoptera: Hesperiidae) of the United States: biology, morphology, and descriptions. Gainesville, University of Florida. PhD dissertation. Núñez-Bustos, E. O. 2008. Diversidad de mariposas diurnas em La Reserva Privada Yacutinga, província de Misiones, Argentina (Lepidoptera: Hesperioidea y Papilionoidea). Tropical Lepidoptera Research 18(2): 7887. Núñez-Bustos, E. O. 2009. Mariposas diurnas (Lepidoptera: Papilionoidea y Hesperioidea) del Parque Nacional Iguazú, província de Misiones, Argentina. Tropical Lepidoptera Research 19(2): 71-81. Orlandin, E., Piovesan, M., Carneiro, E. 2020. Borboletas do Meio-Oeste de Santa Catarina: História Natural e Guia de Identificação. Joaçaba (Brazil), Clube dos Autores. 400 pp. Raby, R., Siewert, R. R., Freitas, A. V. 2024. Immature stages of Rhinthon bajula (Schaus, 1902) (Hesperiidae: Hesperiinae). Tropical Lepidoptera Research 34(2): 118-121. Sahoo, R. K., Warren, A. D., Collins, S. C., Kodandaramaiah, U. 2017. Hostplant change and paleoclimatic events explain diversification shifts in skipper butterflies (Family: Hesperiidae). BCM Evolutionary Biology 17(174): 1-9. Scoble, M. J. 1992. The Lepidoptera: Form, Function and Diversity. New York, Oxford University Press. 404 pp. Schneider, C. A., Rasband, W. S., Eliceiri, K. W. 2012. NIH Image to ImageJ: 25 years of image analysis. Nature Methods 9(7): 671-675. Sierra-Botero, L., Calonje, M., Robbins, R. K., Rosser, N., Pierce, N. E., López-Gallego, C., Valencia-Montoya, W. A. 2022. Cycad phylogeny predicts host plant use of Eumaeus butterflies. Ecology and Evolution 13(4): e09978. Sherer, A., Raghuraman, S., Nakahara, S., Gallice, G. 2025. Immature stages and host plant records for the skipper butterflies Xeniades orchamus (Cramer, 1777), Dubia dubia (Bell, 1932), and Tricrista canta (Evans, 1955) in the Peruvian Amazon (Lepidoptera: Hesperiidae: Hesperiinae: Hesperiini). Zootaxa 5609(3): 390-404. Souza, J. A., Medeiros, A. D., Dolibaina, D. R., Kerpel, S. M. 2023. A new species of Vehilius Godman, 1900 (Lepidoptera, Hesperiidae, Hesperiinae) endemic to the Caatinga, northeastern Brazil: taxonomy, biology and behavior. Zootaxa 5360(2): 239-254. Toussaint, E. F., Braby, M. F., Müller, C. J., Petrie, E. A., Kawahara, A. Y. 2022. Molecular phylogeny, systematics and generic classification of the butterfly subfamily Trapezitinae (Lepidoptera: Papilionoidea: Hesperiidae). Zoological Journal of the Linnean Society 195: 1407-1421. Tscharntke, T., Greiler, H.-J. 1995. Insect communities, grasses, and grasslands. Annual Review of Entomology 40: 535-558. Vega, Y., Flores Bancayan, C., Cordova Velazco, A., Escalante Arteaga, Z., Gallice, G., Nakahara, S. 2025. Immature stages and biology of the Neotropical skipper Artines angelica Medeiros, 2019 (Hesperiidae: Hesperiinae: Hesperiini: Moncina). Acta Amazonica 55: e55bc24162. War, A. R., Paulraj, M. G., Ahmad, T., Buhroo, A. A., Hussain, B., Ignacimuthu, S., Sharma, H. C. 2012. Mechanisms of plant defense against insect herbivores. Plant Signaling & Behavior 7(10): 1036-1320. Warren, A. D., Ogawa, J. R., Brower, A. V. Z. 2009. Revised classification of the family Hesperiidae (Lepidoptera: Hesperioidea) based on combined molecular and morphological data. Systematic Entomology 34: 467-523. Zikán, J. F., Zikán, W. 1968. Inseto-fauna do Itatiaia e da Mantiqueira. Pesquisa Agropecuária Brasileira 3: 45-10. TROP. LEPID. RES., 35(2): 117-121, 2025SUÊNIA-BASTOS ET AL.: Immatures of Orthos hyalinus