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Cladistics of the Ceratocampinae (Lepidoptera: Saturniidae)

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Cladistics of the Ceratocampinae (Lepidoptera: Saturniidae) Vol. 8 Suppl. 2 1997 BALCAZAR and WOLFE: Ceratocampinae 1 TROPICAL LEPIDOPTERA, 8 (Suppl. 2): 1-53 CLADISTICS OF THE CERATOCAMPINAE (LEPIDOPTERA: SATURNIIDAE) MANUEL A. BALCAZAR-LARA AND KIRBY L. WOLFE Depto. de Zoologia, Inst. de Biologfa, Universidad Nacional Autonoma de Mexico, A.P. 70-153, Mexico City, 04510 D.F., Mexico 3090 Cordrey Dr., Escondido, California 92029, USA Contemporary insect systematists must face two distinct gies are used to study the relationships among Saturniidae taxa, challenges. One is the provision of descriptive and analytical and Minet (1994) rehabilitated Draudt's (1927) concept of the taxonomic reports on the insects, producing the cladistic patterns Saturniidae. This conception treats the "Oxytenidae" and "Cerco- that serve as the general frame of reference for comparative phanidae," two "aberrant families" as originally described by entomology and the source of evolutionary patterns in need of Jordan (1924), as two saturniid subfamilies. Further, Minet (1994) causal explanations. The other is to inventory effectively a fauna stated that the Oxyteninae are likely to represent the most that may be 2 to 60 times as diverse as all insects described "primitive" lineage of the Saturniidae, based on antennal charac- during the past 200 years (Wheeler, 1990). ters. The Ceratocampinae is the second most numerous subfamily of Lemaire (1988) and Michener (1952) studied the morphology the New World Saturniidae. With 170 known species, distributed of the adults. Although there is a lack of technical descriptions of in 27 genera, it includes some of the largest and smallest taxa of Saturniidae larvae (e.g., Heppncr and Wang, 1987), the immatures the family. The subfamily is distributed from southern Canada to of some Ceratocampinae have been reared in recent times. The northern Argentina, with a high percentage of species associated present study incorporates available information on the immature with deciduous or partially deciduous dry forests (Lemaire, 1988). stages, both from the literature and from preserved larvae. In general, the peculiar male antennae (quadripectinate basally The objectives of the present study were as follows: and simple distally), the sphingid-like shape of the male, and the 1. Identify monophyletic groups within the Ceratocampinae, using very ornate armature of the early instar larvae, characterize the both adult and larval characters. Ceratocampinae (Michener, 1952; Lemaire, 1988). 2. Provide a comprehensive analysis of the adult and larval The Ceratocampinae have been the subject of three outstanding morphology of Ceratocampinae. modern works. Michener (1952) published a morphological treatment of the Saturniidae, which dealt with genera and higher MATERIAL AND METHODS taxa of the New World, plus subfamilies of the Old World. The Taxa classification proposed by Michener (1952) has been widely used All the genera of the Ceratocampinae were considered as the since by researchers in the field. In his study, Michener included ingroup for the study. Those taxa for which subgenera or "species all the genera of Ceratocampinae (under Citheroniinae) known at groups" have been proposed, were considered as independent the time and proposed several new genera and subgcnera for the entities and coded for accordingly (i.e., Psilopygida (P.) and P. subfamily. Ferguson (1971-72) provided an excellent revision for (Psigida) and the six species groups of Syssphinx proposed by the group in North America. Lemaire (1988) published a revision Lemaire, 1987). These taxa made a total of 33 taxa for the for the whole subfamily for the first time. ingroup (27 genera, two subgcnera and six species groups) (Table There are several aspects of Ceratocampinae that make it an 1). For all the taxa included in the ingroup, at least one species interesting group for a cladistic analysis. It has been considered was thoroughly studied, and literature characters were always a "primitive" group among Saturniidae (Michener, 1952; Lemaire, compared against actual specimens (with the exception of 1988), a condition for considering the subfamily as an outgroup Mielkesia, for which we had no specimens). candidate in future cladistic analyses of other subfamilies. Due to Michener (1952) and Lemaire (1987) concluded that Arsenuri- the extensive revision of Lemaire (1988), there is a sound nae is phylogenetically nearer to Ceratocampinae than to any taxonomic knowledge of the group, an essential requirement for other subfamily of Saturniidae. And some authors, such as Jordan any cladistic analysis. Although it is usually easy to define genera (1922) even united Ceratocampinae and Arsenurinae into by apomorphic characters, the determination of relationships "Arsenuridae", distinct from Saturniidae, but conforming the between genera is far more difficult (Lemaire, 1988), and in supcrfamily Saturnioidea together. Minet (1994) showed that general the higher classification of Saturniidae is still far from Oxytenis Hiibner, Asthenidia Westwood and Homoeopterys Jordan stable (Minet, 1986; Holloway, 1987). Recently, Peigler (1989, form a monophyletic group and, because he concluded that they 1993) published the first two papers in which cladistic methodolo- represent the most "primitive" lineage of the Saturniidae, he TABLE 1- Taxa included in the cladistic analysis (ingroup and outgroup), number of species known, their geographical distribution, references to immature illustrations or descriptions and food plants reported (based mainly on Stone, 1991). Taxa Geographical distribution Larval illustrations or descriptions Hostplants INGROUP Eacles 17 s. Canada to n. Argentina Crocomo & Parra, 1979; Feige, 1971; Aceraceae, Anacardiaceae, Betulaceae, Comaceae, Cupressaccac. Gage, 1976; Lampe, 1986; Leinaire, Ebenaceae, Fabaceae, Fagaceae, Hamamclidaceae, Juglandaceae, 1987; Packard, 1905 Malvaceae, Melastomataceae, Oleaceae, Pinaceae, Platanaceae, Rosaceae, Ulmaceae Bathyphlebia 5 sw. Venezuela to se. Peru (Andean) Citheronia 21 n. USA to n. Argentina Bourquin, 1945; Burmeister, 1879; Anacardiaceae, Aquifoliaceae, Cayocaraceae, Combretaceae, Dias, 1978a, 1981, 1982; Janzen, 1982; Euphorbiaceae, Hamamelidaceae, Juglandaceae, Loranthaceae, Lampc, 1985; Lemaire, 1987; Packard, Malvaceae, Myrtaceae, Oleaceae, Salicaceae 1905; Tuskes, 1987; Vazquez, 1944 Procitheronia 3 French Guiana to Bolivia Dias, 1991 Croton floribundus (Euphorbiaceae) and se. Brazil Schausiella 11 Costa Rica to Bolivia and Lemaire, 1987; Travassos, 1958 Hymenaea (Fabaceae: Caesalpinioideae) se. Brazil Othorene 4 Mexico to se. Brazi Lemaire, 1987; Travassos & Noronha, Terminalia (Combretaceae), Gleditsia (Fabaceae: Caesalpinioideae), 1 1967b Quercus (Fagaceae), Psidium (Myrtaceae), Manilkara (Sapotaceae) Cicia 5 Guiano-Amazonian region Gleditsia (Fabaceae: Caesalpinioideae) and c.-se. Brazil. Dacunju 1 c.-se. Brazil Giacomellia 2 Bolivia and n. Argentina (sub-Andean areas) Aimeidella 3 se. Brazil Otero, 1971 Psilopygoides 1 Guianan-Amazonian Ceropoda 1 Mato Grosso (Brazil) and Paraguay Mielkesia 1 Curiliba (Parana, Brazil) Psilopygida 1 Argentina and e. Bolivia Bourquin, 1945 Gleditsia (Caesalpinioideae), Acacia (Fabaceae: Mimosoideae) (Psilopygida) (Andes) Psilopygida 2 Tropical South America (Psigida) Syssphinx Mexico to Argentina Packard, 1905; Travassos & Ferreira Cassia, Gleditsia (Caesalpinioideae), Robinia (Faboideae), molina gp. d'Almeida, 1937 Acacia, Albizia, Pithecellobium (Fabaceae: Mimosoideae), Salix (Salicaceae) Svssphinx quad- Mexico to Ecuador Heitzman, 1961; Tuskes, 1986 Cassia, Gleditsia, Haematoxylum, Parkinsonia (Caesalpinioideae). rilineata gp. Robinia (Faboideae), Acacia, Albizia, Pithecellobium, Prosopis (Fabaceae: Mimosoideae) Sys sphinx 16 Mexico to Ecuador Comstock, 1947; Tuskes, 1986 Cercidium, Gleditsia, Gymnocladus, Parkinsonia (Caesatpu -oideae), bisecta gp. Robinia (Faboideae), Acacia, Pithecellobium, Prosopis (Faoaceae: Mimosoideae) Syssphinx I se. Canada to ne. Mexico Gleditsia, Gymnocladus, Hymenaea (Caesalpinioideae), Robinia bicolor gp. (Fabaceae: Faboideae). Quercus (Fagaceae) Svssphinx I Amazonian amena gp. Syssphinx 3 e. slopes of Andes, n. Venezuela to Bolivia ocellata gp. Anisota 13 Canada to Guatemala Riottc & Peigler, 1980; Wolfe & Acer (Aceraceae), Corylus (Betulaceae) Castanea, Fagus, Peigler, 1993 Quercus (Fagaceae), Carya (Juglandaceae), Rubus (Rosaceae) Dryocampa 1 sc. Canada to Florida and Allen, 1976 Acer (Aceraceae), Fagus, Quercus (Fagaceae), Juglans (Juglandaceae) e. Texas Adeloneivaia 15 Mexico to n. Argentina Janzen, 1982; Lampe, 1987; Lemaire, Acacia, Inga, Lysiloma (Fabaceae: Mimosoideae), Quercus 1987; Packard, 1905 (Fagaceae), Salix (Salicaceae) Adelowalkeria 6 Tropical South America Dias, 1986; Lemaire, 1987 Machaerium (Fabaceae : Faboideae) Menace re sa 1 c. Brazil Rachesa 4 Colombia to Bolivia (Andes) Ceratesa 1 e. Bolivia and nw. Argentina Scolesa 6 c.-e. and se. Brazil, nw. Argentina and se. Peru Citioica 2 Mexico to Bolivia Lemaire, 1987 Robinia (Fabaceae: Faboideae), Salix (Salicaceae) Oiticella 3 Venezuela to se. Brazil and Argentina Mimosa (?) (Fabaceae : Mimosoideae) Pliloscoia 9 Mexico to Bolivia and Janzen, 1982; Lemaire, 1987 Acacia (Fabaceae: Mimosoideae) se. Brazil Neorcarnegia 1 c.-se. Brazil to e. Bolivia OUTGROUP Arsenura 20 Mexico to Bolivia and Brenner & Lampe, 1987 Annona (Annonaceae),Bwn/w«.v>/WJ,s, Bombax, Chorisia (Bombacaceae), s. Brazil Urena (Malvaceae), Tilia (Tiliaceae) Dysdacmonia 3 Mexico to Bolivia and Dias, 1978b Bombax, Ceiba, Chorisia (Bombacaceae), Malvaceae, Tiliaceae
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