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 MORAES
1,2
* and MARCELO DUARTE
2
1
Curso 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
2
Museu 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
doi: 10.1111/zoj.12102
ADDITIONAL KEYWORDS: Batesian mimicry – new synonyms – taxonomy.
INTRODUCTION
Currently Castniidae is placed in the superfamily
Cossoidea (van Nieukerken et al., 2011). Although a
molecular analysis showed a phylogenetic affinity
between Castniidae and Cossidae, it also showed
phylogenetic affinities between Castniidae and
Sesiidae (Regier et al., 2009). Indeed, Edwards et al.
(1998) placed Castniidae together with Sesiidae and
Brachodidae, based on adult characters provided by
Minet (1991) and Kozlov, Kuznetzov & Stekolnikov
(1998). Castniidae is distinguished by the presence of
a ridge on the prothoracic tergite, the saccus with
anterior projections, the ejaculatory bulb enlarged,
the extendable ovipositor, and the hooked aedeagus
(Edwards et al., 1998); however, these features are
also found in other families of Lepidoptera.
According to Miller (1986), Castniidae also has
some similarities with Tortricidae and Cossidae, as
previously speculated based on some characters of
immatures, such as vertical eggs, larvae with the
borer habit, and pupae with spines on the abdominal
terga (Mosher, 1916). Nevertheless, castniids
differ from the Tortricidae by the absence of
chaetosemata, and from the Cossidae by the presence
of an apiculus on the distal part of the antenna
(Miller, 1986).
The family Castniidae contains approximately 150
species, and includes the subfamilies Castniinae
*Corresponding author. E-mail: simeao_moraes@yahoo.
com.br
Zoological Journal of the Linnean Society, 2014. With 143 figures
© 2014 The Linnean Society of London, Zoological Journal of the Linnean Society, 2014 1