Phylogenetic relationships of butterflies of the tribe Acraeini ...

526 K.L. Silva-Brandão et al. / Molecular Phylogenetics and Evolution 46 (2008) 515–531genus Pardopsis also appeared within the Argynnini cladewith a high PP value.3.2. Evolution of host plant useAccording to the character optimization of the hostplant families used by each species of Acraeini, the ancestralstate of the tribe is the use of host plants of the familyPassifloraceae, and most of the basal Acraea + Bematistesfeed on plants of this family (Fig. 5). The species of Acraeaof this basal clade also feed on Violaceae. The clade joiningthe remaining Acraea (‘‘Old World Actinote”) is characterizedby feeding mostly on Urticaceae, with some specieshighly polyphagous—the branch appears generally as‘‘equivocal” to this polyphagous species. The host plantused by the ancestor of the South-American Acraeini cladeis equivocal, but if we consider the whole clade but Alt. eresia(whose host plant is unknown) the ancestor host plantof this group may have been Asteraceae, and all NeotropicalAcraeini with a well known life cycle feed on plants ofthis family only. Feeding on this host plant family is characteristicof the ancestor of the genus Actinote (s. str.). Thephylogenetic signal obtained by the modified PTP test wassignificant (P = 0.003), suggesting that the distribution ofthis trait among taxa can be explained by their phylogeneticrelationships.4. Discussion4.1. The internal relationships of HeliconiinaeAlthough the main subject of this study has been therelationships within the tribe Acraeini, it is worth discussingthe position of the tribe within the subfamily Heliconiinae.Even with all the recent advances in the systematics ofNymphalidae, the affinities of the tribe Acraeini within theHeliconiinae continue to be undefined and controversial.The recent phylogenetic proposal of Penz and Peggie(2003) for Heliconiinae, based on characters of early stagesand adult morphology, showed a polytomy among the fourmajor clades in the trees obtained with equally weightedcharacters; however, the successive approximation weightinganalyses resulted in a better resolved tree, with theAcraeini appearing as the sister group of Heliconiini+ (Vagrantini + Argynnini). The same results werefound by Freitas and Brown (2004) with independent datasets of morphological characters from adults and immatures.Alternative hypotheses were proposed by Brower(2000) and Wahlberg et al. (2003b, 2005), with the tribeArgynnini appearing as sister to Acraeini + Heliconiini.The taxonomic sampling within Heliconiinae in these studies,however, is too limited (usually one or two taxa representingeach tribe) to permit further discussion on the tribalrelationships. Here we found two different results in relationto the tribal relationships within Heliconiinae,although our sample does not include many representativesof other tribes outside Acraeini. The results achieved by theBayesian phylogenetic hypothesis are similar to that foundby other studies, with Vagrantini + Argynnini (containingPardopsis) sister to Heliconiini (+Cethosiini) + Acraeini.On the other hand, the MP consensus tree shows the tribeHeliconiini sister to the other three tribes.The monophyly of the tribe Acraeini was recovered inall results of Penz and Peggie (2003) and Freitas and Brown(2004) (with Pardopsis as part of Acraeini in both studies),and the present study has also recovered the monophyly ofthe tribe Acraeini (s. str.) in all MP and Bayesian analyses,although all results have shown that Pardopsis does notbelong to Acraeini. The monotypic genus Pardopsis hasbeen considered as part of Acraeini and sister to all othergenera of this tribe (Penz and Peggie, 2003; Freitas andBrown, 2004). However, this taxon was never included ina molecular study before, and the results of the presentanalyses showed that the genus is in fact part of Argynnini.The position of Pardopsis within Argynnini was recoveredby the Bayesian analysis with a high posterior probabilityvalue (Fig. 4), and in the consensus MP tree with a weaksupport, and we consider it not to be within the tribeAcraeini (s. str.), as defined in this study. Some clues aboutthe position of Pardopsis can be obtained from the immaturestages (see Van Son, 1963), which share several characterstates with the Argynnini, as discussed in Freitas (1999).These character states include, for example, the egg withseveral longitudinal ridges anastomosing near half theheight, and the subdorsal prothoracic scoli longer thanthe others (both present in Euptoieta hegesia—Freitas,1999). A future study with better taxonomic sampling ofArgynnini and Vagrantini, and the addition of morphologicaldata, will be crucial to define the real affinities of thispuzzling genus.The position of Cethosia within the Heliconiinae hasbeen discussed in several recent papers, but still remainscontroversial (Penz and Peggie, 2003). Brower (2000)shows Cethosia as sister to Acraeini + Heliconiini. In Freitasand Brown (2004), Cethosia is the sister group of Vindula(=Vagrantini of Penz and Peggie, 2003), and thesuccessive weighting analysis defined this pair as sister toHeliconiini. The same general topology was also foundby Penz and Peggie (2003), and the Bayesian analysis inthe present study shows Cethosia as sister to Heliconiini(Fig. 4). Harvey (1991), however, proposes that Cethosiabelongs to its own tribe, Cethosiini, sister to the Acraeini,based on an extremely well developed ‘‘gland sous-papillaire”in the female abdomen, although in the ‘‘Addendum”of the same work the author places Cethosia intoAcraeini and abandons Cethosiini. Clearly a broad molecularstudy of the subfamily Heliconiinae is needed with acomprehensive sampling of species in the subfamily to clarifythe position of Cethosia.4.2. The tribe AcraeiniOur results on the relationships within Acraeini are similarto the proposal of Pierre (1987) (Fig. 2), but due to the