Although mutualisms have been intensively investigated, demonstration of indirect effects between co-occurring mutualistic systems is rare. For instance, the ecological consequences of co-occurrence of ant-tended insects on a plant have never been examined for survival effects on either trophobiont species. Here, we assess the selective pressures mediating co-occurrence of a facultative ant-tended butterfly (Parrhasius polibetes) with ant-tended treehoppers (Guayaquila xiphias) on Schefflera vinosa shrubs. We evaluated host plant selection and caterpillar survival in P. polibetes in the presence and absence of ant-treehopper associations. Paired trials revealed that butterflies preferably oviposit on branches hosting ant-tended treehoppers when they had a choice between those and branches without this interaction. Presence of ant-tended treehoppers on a branch reduced the abundance of P. polibetes' natural enemies and improved caterpillar survival in both premyrmecophylic and ant-tended phases. Thus ant-tended treehoppers create an enemy-free space on foliage that butterflies exploit to protect larval offspring. These findings connect two widely documented ant-trophobiont mutualisms and highlight the importance of considering multiple interactions for a proper understanding of ant-plant-herbivore systems. Detection of other ant-based mutualisms on oviposition to improve offspring survival may have represented an important evolutionary step in the process of host plant selection in facultative myrmecophilous butterflies.

ABSTRACT. External morphology of the immature stages of neotropical heliconians: IV. Dryas iulia alcionea (Lepidoptera, Nymphalidae, Heliconiinae). The external features of egg, larva and pupa of Dryas iulia alcionea (Cramer, 1779) are described and illustrated.

The underlying mechanisms mediating the use of multiple host plants were investigated in Parrhasius polibetes (Lycaenidae), a florivorous and facultative myrmecophilous butterfly. Plant traits such as presence of ant-treehopper associations as a source of enemy-free space, flower bud dimensions, toughness, thickness, trichomes, and the corresponding performance and wear of P. polibetes mandibles were examined for three natural hosts: Schefflera vinosa (Araliaceae), Pyrostegia venusta (Bignoniaceae) and Luehea grandiflora (Malvaceae). Parasitism levels of larvae found on the three hosts were also determined. Almost all Luehea had ant-treehopper associations, and all larvae found on this host were non-parasitized. Parasitism was low in larvae found on Schefflera, half of which hosted ant-treehopper associations. No ant-treehopper association was found on Pyrostegia, where parasitism was significantly higher compared to other hosts. In the laboratory, P. polibetes performed well on Schefflera, followed by Pyrostegia. No larvae survived when fed with Luehea. Flower buds of Luehea were thicker and tougher than those of Schefflera and Pyrostegia. Indeed, mandibles of larvae reared on Luehea showed substantial wear, whereas those reared either on Schefflera or Pyrostegia presented no significant damage. Additionally, we suggest that co-occurrence with ant-treehopper associations on a plant provides parasitoid-free space for P. polibetes larvae. Our results support the hypothesis that ecological trade-offs among host plants (i.e., food quality and enemy-free space) promote polyphagy in natural populations of P. polibetes. Host morphological traits seem to play a relevant role in P. polibetes performance. To our knowledge, this is the first report showing the costs of polyphagy in a myrmecophilous butterfly.

ABSTRACT. External morphology of the immature stages of neotropical heliconians: V. Agraulis vanillae maculosa (Lepidoptera, Nymphalidae, Heliconiinae). The external features of egg, larva and pupa of Agraulis vanillae maculosa (Stichel, [1908]) are described and illustrated, based upon light and scanning electron microscopy.

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ABSTRACT. Externai rnorphology of the irnrnature stages of neotropical heliconians. IH. Heliconills erato phyllis (Fabricius) (Lepidoptera, Nymphalidae, Heliconiinae). The externai features of egg. larva and pupa of Heliconius erato phyllis (Fabricius, 1775) are described and illustrated, based upon light and scanning electron microscopy. KEY WORDS. Lepidoptera, Heliconiini , Nymphalidae, immalure stages, morphology He/iconius Kluk, 1802 (Lepidoptera, Nymphalidae) é um dos gêneros mais conspícuos e co nhecidos de borbo letas neotropicais (DEVRIES 1987). Heliconius erato (Li nnaeus , 1758) é considerada a espéc ie mai s vari ável do gênero, sendo reconhec id as, até o presente, 29 subespéc ies (HOLZINGER & HOLZINGER 1994). Segund o os mesmos autores, He/iconius erato phyllis (Fabricius, 1775) apresenta ampla distribuição na América do Su l, ond e ocorre desde o nord es te até o sul do Brasil , Bolívia, nordeste da Argentina e Uruguai.Heliconius erato é comumente encontrada em ambi entes enso larados, sendo freqüente em florestas abertas ou perturbadas, bem como clareiras em florestas den sas (BROWN 1992). Os adultos possuem comportamento semigregá-rio , fo rmand o agrupamen tos noturnos chamados de "dormi tórios " que reúnem de 2 a 145 indi víduos (CRANE 1957; TURNER 1971;MALLET 1986 ; R. DiMare, UFRGS , dados não pub li cados) . O ovo é depositado na região apical dos ramos de várias espécies de Passiflora Linnaeus, preferencialmente do sub gê nero Plectostemma Masters. A larva é so litária e apresenta tendência ao canibali smo (B ROWN & MIELKE 1972; BENSON et ai. 1976). As espécies de Heliconius , dev ido à fác il adaptação às cond ições de insetário e cu lti vo de suas plantas hospedeiras, têm si do utili zadas como modelos experim enta is em diversos es tudos so bre interação in seto-planta

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Riodinidae is a highly diverse butterfly family with the majority of its genera restricted to the Neotropics and, despite previous efforts, its higher systematics remains unresolved. Here, we propose a novel phylogenetic hypothesis, based on a comprehensive sample of riodinids, primarily from the Neotropics, covering 67% of all genera and all of the major lineages. We sequenced nine molecular markers and estimated resulting phylogenies with maximum likelihood and Bayesian approaches, using both timed trees and time-independent trees. We based calibration on three fossil Riodinidae, and reassessed the position of the oldest fossil. We also incorporated 52 samples from a previous study providing a comprehensive maximum likelihood tree for 304 species comprising 80% of all genera. We propose a new higher classification of the Riodinidae with two subfamilies