bs_bs_banner Zoological Journal of the Linnean Society, 2014, 170, 362–399. With 143 figures Phylogeny of Neotropical Castniinae (Lepidoptera: Cossoidea: Castniidae): testing the hypothesis of the mimics as a monophyletic group and implications for the arrangement of the genera SIMEÃO DE SOUZA MORAES1,2* and MARCELO DUARTE2 1Curso de Pós-Graduação em Ciências Biológicas (Zoologia), Instituto de Biociências, Departamento de Zoologia, Universidade de São Paulo, Rua do Matão, travessa 14, número 321, CEP 05508-900, São Paulo, São Paulo, Brazil 2Museu de Zoologia da Universidade de São Paulo, Avenida Nazaré 481, 04263-000, São Paulo, São Paulo, Brazil Received 19 March 2013; revised 11 October 2013; accepted for publication 13 October 2013 A cladistic analysis of the Neotropical Castniidae is presented using 120 morphological characters, and a taxonomic treatment based on that analysis is also presented. The tribe Gazerini as previously delimited was found to be paraphyletic with respect to the genera Ceretes, Divana, Riechia, Frostetola, and Oiticicastnia. The genera Castnia, Geyeria, and Athis were also found to be non-monophyletic taxa. The mimicry pattern had multiple independent origins in the Neotropical castniids, and at least two lineages, Riechia and Prometheus, are involved in Batesian mimicry rings with unpalatable butterfly models in the tribes Acraeini and Heliconiini (Nymphalidae). We propose for Castniini 13 new synonymies and 27 new combinations. Geyeria strigata (Walker, 1854) is revalidated. The generic placements of Athis superba (Strand, 1912) and Castnia eudesmia Gray, 1838 are questionable, but presently upheld. © 2014 The Linnean Society of London, Zoological Journal of the Linnean Society, 2014, 170, 362–399. doi: 10.1111/zoj.12102 ADDITIONAL KEYWORDS: Batesian mimicry – new synonyms – taxonomy. INTRODUCTION anterior projections, the ejaculatory bulb enlarged, the extendable ovipositor, and the hooked aedeagus Currently Castniidae is placed in the superfamily (Edwards et al., 1998); however, these features are Cossoidea (van Nieukerken et al., 2011). Although a also found in other families of Lepidoptera. molecular analysis showed a phylogenetic affinity According to Miller (1986), Castniidae also has between Castniidae and Cossidae, it also showed some similarities with Tortricidae and Cossidae, as phylogenetic affinities between Castniidae and previously speculated based on some characters of Sesiidae (Regier et al., 2009). Indeed, Edwards et al. immatures, such as vertical eggs, larvae with the borer (1998) placed Castniidae together with Sesiidae and habit, and pupae with spines on the abdominal terga Brachodidae, based on adult characters provided by (Mosher, 1916). Nevertheless, castniids differ from the Minet (1991) and Kozlov, Kuznetzov & Stekolnikov Tortricidae by the absence of chaetosemata, and from (1998). Castniidae is distinguished by the presence of the Cossidae by the presence of an apiculus on the a ridge on the prothoracic tergite, the saccus with distal part of the antenna (Miller, 1986). The family Castniidae contains approximately 150 *Corresponding author. E-mail: simeao_moraes@yahoo. species, and includes the subfamilies Castniinae com.br and Tascininae (Edwards et al., 1998). Studies on 362 © 2014 The Linnean Society of London, Zoological Journal of the Linnean Society, 2014, 170, 362–399 PHYLOGENY OF NEOTROPICAL CASTNIINAE 363 Tascininae are few, as are the number of specimens cal properties, or morphological features of the held in collections worldwide. Currently the group castniids with a view towards understanding the comprises four described species belonging to the relationships among the species and the evolution genus Tascina Westwood, 1877, which occur in the of this group. Here, we provide a morphological Indo-Malayan region (Holloway, 1998; Fukuda, 2000). data set based on characters other than colour, in Castniinae is a more diverse group, arranged in three order to test the monophyly of Neotropical tribes and tribes. Synemoniini, found in the Australian region, is relationships among the genera. Based on this pool represented by 24 described species. of morphological data, we construct a phylogenetic The other two tribes of Castniinae (Gazerini and hypothesis, and suggest new rearrangements at the Castniini) occur in the Neotropics and comprise 31 genus level for the Neotropical castniids, based on genera and 88 species; together these are the most monophyletic groups. We also provide an overview diverse and speciose group of Castniidae (Figs 1–36). of the taxonomy of Castniidae, focusing on the The two tribes are inadequately delimited, however, Neotropical species. We briefly discuss the evolution based only on colour pattern. Gazerini includes the of the colour patterns that mimick certain butter- mimetic species of Ithomiinae and Heliconiinae flies, and hope that this introductory analysis may (Nymphalidae), and the remaining species are encourage further studies focusing on different included in Castniini (Lamas, 1995; Miller, 1995). aspects of the mimicry, by alternative methods (e.g. The only phylogenetic analysis that tested the DNA sequencing). monophyly of Castniidae and the relationships among the subfamilies, tribes, and genera was performed by Miller (1986). In this study, Castniidae and its three A TAXONOMIC OVERVIEW OF THE constituent subfamilies were all recovered as a NEOTROPICAL SPECIES monophyletic group, which is accepted as the starting point for the present investigation; the Neotropical The first record of a castniid was an illustration of an Castniinae + Synemoniini formed the sister group of unknown species by Maria Sibylla Merian (1705), the Tascininae. Miller (1986) also recovered the Neo- later described as Papilio licus by Drury (1773). tropical Castniini and Gazerini as monophyletic taxa. Fabricius (1807) erected 40 genera of Lepidoptera Although some genera have been synonymized from the genus Papilio, the seventh of which was (González & Cock, 2004; Moraes & Duarte, 2009), Castnia Fabricius, 1807. In 1824, Latreille & Godart some monotypic genera of the Neotropical Castniidae and Dalman published the first studies compiling are still considered valid (Table 1), based on wing species of castniids, with both works focusing on colour pattern, without further support from morpho- colour characters. Latreille & Godart (1824) gave a logical evidence. For more than a century the brief redescription of eight species, most of which Neotropical Castniidae have received typological– were originally described by Cramer (1773–1782) and taxonomic treatments, mainly based on colour Fabricius (1775), and described five new species. pattern, and some species that are presently consid- Dalman (1824) dealt with 18 species, of which four ered as valid may actually represent chromatic vari- were considered new. Some of the species treated by ations, as shown by Moraes, Duarte & González Latreille & Godart (1824) were not mentioned by (2010), and suggested by Vinciguerra (2011b). Dalman (1824), and vice versa. Gray (1838) published In the Neotropics, some castniids are economically a synopsis of Castnia including 29 valid species, three important agricultural pests that cause extensive of which were new. Although the species were still damage to plantations of sugar cane, oil palms, pine- grouped within the genus Castnia, the author pro- apples, and orchids. They have been studied with posed five sections, using characters such as the regard to pest control (Esquivel, 1981), chemical colour and shape of wings, proportions of palp seg- (Rebouças, Caraciolo & Sant’Ana, 1999), morphologi- ments, and presence or absence of sexual dimor- cal (Moraes & Duarte, 2009), ethological (Albertoni phism. He also discussed differences in the branching et al., 2012), and molecular aspects (Silva-Brandão pattern of veins of the radial system. Between 1854 et al., 2012). The main agricultural pest species is and 1858, Walker (1854), Ménétriés (1857), and Telchin licus (Drury, 1773), which bores into sugar- Herrich-Schäffer (1850–1858) provided colour-based cane stem tissues. Other important pest species are descriptive studies. Walker attempted to separate Eupalamides cyparrissias (Fabricius, 1776) on palm the 41 species described or redescribed in his study trees, Riechia acraeoides (Guérin-Ménéville, [1832]) into subdivisions, using the colour and shape of the on orchids, and Castnia invaria Walker, 1854 on wings, but provided no further discussion on tax- pineapples. onomy. In the study by Boisduval (1875), Castniidae Few studies have investigated the butterfly-like was represented by 78 species divided into the tribes behaviour (Sarto i Monteys et al., 2012), physiologi- ‘Castniaires’ and ‘Synemonides’. Synemonides was © 2014 The Linnean Society of London, Zoological Journal of the Linnean Society, 2014, 170, 362–399 364 S. S. MORAES AND M. DUARTE © 2014 The Linnean Society of London, Zoological Journal of the Linnean Society, 2014, 170, 362–399 PHYLOGENY OF NEOTROPICAL CASTNIINAE 365 Table 1. Genus arrangement for the Neotropical Castniidae according to Lamas (1995) and number of species per genus included in the cladistic analysis Number of Number Number of Number species per of species species per of species Genera genus* dissected Genera genus* dissected Athis Hübner, [1819] 16 6 Yagra Oiticica, 1955 2 2 Castnia Fabricius, 1807 6 2 Feschaeria Oiticica, 1955 2 1 Eupalamides Hübner, [1819] 5 1 Tosxampila Oiticica, 1955 2 1 Synpalamides Hübner, [1823] 5 2 Ceretes Schaufuss,