Lepidoptera: Nymphalidae: Satyrinae), a Rare and Endangered Butterfly from Southeastern Brazil
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Neotrop Entomol (2012) 41:461–467 DOI 10.1007/s13744-012-0073-5 SYSTEMATICS, MORPHOLOGY AND PHYSIOLOGY Euptychia boulleti (Le Cerf) n. comb. (Lepidoptera: Nymphalidae: Satyrinae), a Rare and Endangered Butterfly from Southeastern Brazil 1 2 2 3 4 AVL FREITAS ,NWAHLBERG ,PFMATOS-MARAVI ,MAMARIN , OHH MIELKE 1Depto de Biologia Animal, Museu de Zoologia, Instituto de Biologia, Univ Estadual de Campinas, Campinas, SP, Brasil 2Lab of Genetics, Dept of Biology, Univ of Turku, Turku, Finland 3Grupo de Investigación en Sistemática Molecular, Univ Nacional de Colombia, Medellín, Colombia 4Depto de Zoologia, Univ Federal do Paraná, Curitiba, PR, Brasil Keywords Abstract Atlantic forest, Caenoptychia, Euptychiina, This paper discusses the systematic position of the rare and endangered morphology satyrine butterfly Caenoptychia boulleti Le Cerf, the only included species Correspondence in Caenoptychia (type species), based on adult morphology and molecular AVL Freitas, Depto de Biologia Animal, data. The results showed that Caenoptychia Le Cerf belongs to the Eupty- Museu de Zoologia, Instituto de Biologia, Univ Estadual de Campinas, 6109, 13083- chia Hübner clade, and the genus is synonymized with Euptychia,new 970 Campinas, São Paulo, Brasil; synonymy. Euptychia boulleti (Le Cerf) is a new combination. The male [email protected] genitalia of E. boulleti showed at least one important synapomorphy with Edited by Roberto A Zucchi – ESALQ/USP the other species of Euptychia, which is the presence of a posterior projection of the tegumen above the uncus. Molecular data reinforces Received 10 September 2011 and accepted the position of Caenoptychia within the genus Euptychia. 24 June 2012 Published online 31 August 2012 * Sociedade Entomológica do Brasil 2012 Introduction Atlantic Forest in Southeastern Brazil, from Espírito Santo to São Paulo (Freitas & Brown 2008). The species is considered Satyrinae (Nymphalidae) is one of the most diverse groups endangered, and the main threats are the destruction and of butterflies, with more than 2,500 species occurring on degradation of its habitats in recent years (Freitas & Brown all continents except Antarctica (Ackery et al 1998, Marín et 2008, Freitas 2010). The ventral wing pattern of this species al 2011). The group is particularly diverse in the Neotropical is so distinctive from all other known satyrines that it was region where the species rich subtribe Euptychiina is rep- compared with some members of the family Lycaenidae by resented by over 400 described species (Lamas 2004). Le Cerf (1919) in its original description. This unique wing This clade, however, is one of the least known groups of pattern probably made this species very conspicuous in early Satyrinae concerning its biology, ecology, and systematics butterfly collections, resulting in three independent descrip- (Marín et al 2011). There are few phylogenetic studies on tions of this taxon in the first half of the twentieth century (Le this group, a large number of undescribed species and Cerf 1919,Joicey&Talbot1924, Gagarin 1936). several polyphyletic genera waiting to be revised (Viloria Since its description, the systematic position of C. boul- 2003, Murray & Prowell 2005, Peña et al 2010, Marín et leti remained enigmatic, and this taxon has not been trea- al 2011). Additionally, there are a number of genera that ted either in the classic morphological work of Miller (1968) are still enigmatic in relation to their affinities within nor in the recent molecular phylogeny of Peña et al (2010). Satyrinae, including the monotypic genus Caenoptychia In the present paper, the systematic position of C. boulleti is Le Cerf. discussed based on morphological and molecular characters, as Caenoptychia boulleti Le Cerf (Fig 1) is an odd and rare well as the taxonomic implications of its position within the species of Euptychiina known from a few sites of montane Euptychiina. 462 Freitas et al Fig 1 Euptychia boulleti, ventral on the left and dorsal on the right. Left, male from Independência, Petrópolis, Rio de Janeiro (lectotype of Ristia tigrina); DZUP 615. Right, female from Maromba, Parque Nacional de Itatiaia, Rio de Janeiro; ZUEC. Material and Methods 10.0×106 generations, sampling trees every 1,000 cycles. The first 4,000 trees were discarded as “burn in” based on Specimens of Caenoptychia boulleti were studied in six when the runs had converged and reached equilibri- collections. The acronyms for the collections are: DZUP, um. Additionally, Bayesian analyses were performed “Departamento de Zoologia, Universidade Federal do Par- on separate gene datasets under the same settings as aná, Curitiba, Paraná, Brazil”; DZUP-OM, “Coleção Olaf H. described above. H. Mielke, Curitiba, Paraná, Brazil”; MNHN, “Muséum Na- tional d’Histoire Naturelle, Paris, France”; MNRJ, “Museu Nacional da Universidade Federal do Rio de Janeiro, Rio de Results Janeiro, Rio de Janeiro, Brazil”; MZSP, “Museu de Zoologia, Universidade de São Paulo, São Paulo, São Paulo, Brazil”; Euptychia Hübner USNM, “National Museum of Natural History, Smithsonian Institution, Washington, DC, USA”; ZUEC, “Museu de Zoo- Euptychia Hübner (1818,p.20).Type-speciesEuptychia logia da Universidade Estadual de Campinas, Unicamp, mollina Hübner (1818), designated by Hemming (1937, Campinas, São Paulo, Brazil” (Table 1). p. 150). Dissections were made using standard techniques. The Caenoptychia Le Cerf (1919, p. 328). Type species C. abdomens were soaked in hot 10% KOH for 10 min before boulleti by original designation. Syn. n. dissection, and dissected parts were stored in glycerol. Taxonomic nomenclature follows Lamas (2004), modified Euptychia boulleti (Le Cerf) n. comb. (Figs 1, 2, and 3) after Peña et al (2006). DNA sequences of cytochrome c oxidase subunit I, Caenoptychia boulleti Le Cerf (1919, p. 328). Holotype male, elongation factor-1 alpha, glyceraldhyde-3-phosphate São Leopoldo, Rio Grande do Sul, Brazil, October, 1905; dehydrogenase, ribosomal protein S5, and wingless MNHN (examined). were sequenced for one individual (4,447 bp total; Euptychia virgata Joicey & Talbot (1924, p. 570). One male DNA voucher PM17-01), as described in Wahlberg & (Holotype), Leopoldina, (unknown state), Brazil; BMNH. Wheat (2008) (GenBank accession numbers: JQ639284, Ristia tigrina Gagarin (1936,p.8),Figs2 and 3. Holotype JQ639285, JQ639286, JQ639287, and JQ639288). (not designated in the collection) and specimens from Octo- These sequences were analyzed together with the ber and November 1934, Independência, Petrópolis, Rio de dataset compiled by Peña et al (2010) and included Janeiro State, Brazil. Lectotype male (designed by Mielke & sequences of Euptychia species from Murray & Prowell Casagrande 1986), Independência, Petrópolis, Brazil, 8 No- (2005), using Bayesian Inference. Bayesian analyses vember 1934; DZUP. (Huelsenbeck et al 2001,Huelsenbecket al 2002)were carried out for the combined data set under the model Diagnosis GTR + Γ, using the program MrBayes 3.2 (Ronquist & Huelsenbeck 2003). Analysis of combined data by Body predominantly brown—wings rounded, brown dor- Bayesian methods permits partition-specific substitution sally (females usually present some yellow discal patches models and parameters (Nylander et al 2004). For between main veins and with a unique ventral pattern of that reason, all substitution model parameters (gamma black stripes pattern over a light yellow background, with a shape, character state frequencies, and substitution marginal white stripe bordered with black. In both sexes, rates of GTR model) were allowed to vary across par- ventral forewing with a patch of orange scales in basal SC titions (0genes). Two simultaneous chains were run for vein, and ventral hindwing with a patch of orange scales in Euptychia boulleti n. comb. a Rare and Endangered Butterfly 463 Table 1 Data for studied individuals of Euptychia boulleti from seven Museum collections (see text for museum acronyms) and field observations. Code Sex Date State Municipality Site Altitude (m) DZUP 1 male 13 Sept 1928 Rio de Janeiro Petrópolis Independência 900 DZUP 1 male 17 Mar 1935 Rio de Janeiro Petrópolis Independência 900 DZUP 1 male 5 Nov 1936 Rio de Janeiro Petrópolis Independência 900 DZUP 1 malea 15 Nov 1936 Rio de Janeiro Petrópolis Independência 900 DZUP 3 males 15 Nov 1936 Rio de Janeiro Petrópolis Independência 900 DZUP 1 male 17 Nov 1936 Rio de Janeiro Petrópolis Independência 900 DZUP 1 femalea 29 Nov 1936 Rio de Janeiro Petrópolis Independência 900 DZUP 2 males 30 Nov 1936 Rio de Janeiro Petrópolis Independência 900 DZUP 1 male 26 Feb 1937 Rio de Janeiro Petrópolis Independência 900 DZUP 1 male 28 Feb 1937 Rio de Janeiro Petrópolis Independência 900 DZUP 1 male 4 Mar 1939 Rio de Janeiro Petrópolis Independência 900 DZUP 1 male 20 Mar 1939 Rio de Janeiro Petrópolis Independência 900 DZUP 1 male 21 Mar 1939 Rio de Janeiro Petrópolis Independência 900 DZUP 1 male 18 Oct 1939 Rio de Janeiro Petrópolis Independência 900 DZUP 1 malea 19 Oct 1939 Rio de Janeiro Petrópolis Independência 900 DZUP 1 male 20 Oct 1939 Rio de Janeiro Petrópolis Independência 900 DZUP 1 male 19 Nov 1939 Rio de Janeiro Petrópolis Independência 900 DZUP 1 malea 1 Jan 1960 Rio de Janeiro Petrópolis Parque São Vicente 930 DZUP 1 male 16 Nov 1960 Rio de Janeiro Petrópolis Independência 900 DZUP 1 femalea 21 Jan 1966 Rio de Janeiro Petrópolis – 1,100 DZUP 1 malea 11 Apr 1968 Rio de Janeiro Petrópolis – 1,200 DZUP-OM 1 male 31 Oct 1964 Rio de Janeiro Itatiaia PNI-Maromba 1,100 DZUP-OM 1 male, 1 female 8 Dec 1936 Rio de