Phylogenetic relationships among major clades of butterflies and skippers have long been controversial, with no general consensus even today. Such lack of resolution is a substantial impediment to using the otherwise well studied butterflies as a model group in biology. Here we report the results of a combined analysis of DNA sequences from three genes and a morphological data matrix for 57 taxa (3258 characters, 1290 parsimony informative) representing all major lineages from the three putative butterfly super-families (Hedyloidea, Hesperioidea and Papilionoidea), plus out-groups representing other ditrysian Lepidoptera families. Recently, the utility of morphological data as a source of phylogenetic evidence has been debated. We present the first well supported phylogenetic hypothesis for the butterflies and skippers based on a total-evidence analysis of both traditional morphological characters and new molecular characters from three gene regions (COI, EF-1alpha and wingless). All four data partitions show substantial hidden support for the deeper nodes, which emerges only in a combined analysis in which the addition of morphological data plays a crucial role. With the exception of Nymphalidae, the traditionally recognized families are found to be strongly supported monophyletic clades with the following relationships: (Hesperiidae+(Papilionidae+(Pieridae+(Nymphalidae+(Lycaenidae+Riodinidae))))). Nymphalidae is recovered as a monophyletic clade but this clade does not have strong support. Lycaenidae and Riodinidae are sister groups with strong support and we suggest that the latter be given family rank. The position of Pieridae as the sister taxon to nymphalids, lycaenids and riodinids is supported by morphology and the EF-1alpha data but conflicted by the COI and wingless data. Hedylidae are more likely to be related to butterflies and skippers than geometrid moths and appear to be the sister group to Papilionoidea+Hesperioidea.

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The need for robust estimates of times of divergence is essential for downstream analyses, yet assessing this robustness is still rare. We generated a time-calibrated genus-level phylogeny of butterflies (Papilionoidea), including 994 taxa, up to 10 gene fragments and an unprecedented set of 12 fossils and 10 host-plant node calibration points. We compared marginal priors and posterior distributions to assess the relative importance of the former on the latter. This approach revealed a strong influence of the set of priors on the root age but for most calibrated nodes posterior distributions shifted from the marginal prior, indicating significant information in the molecular data set. Using a very conservative approach we estimated an origin of butterflies at 107.6 Ma, approximately equivalent to the latest Early Cretaceous, with a credibility interval ranging from 89.5 Ma (mid Late Cretaceous) to 129.5 Ma (mid Early Cretaceous). In addition, we tested the effects of changing fossil calibration priors, tree prior, different sets of calibrations and different sampling fractions but our estimate remained robust to these alternative assumptions. With 994 genera, this tree provides a comprehensive source of secondary calibrations for studies on butterflies.

J. 2004. Diversity and community composition of butterflies and odonates in an ENSO-induced fire affected habitat mosaic: a case study from East Kalimantan, Indonesia. Á/ Oikos 105: 426 Á/446.Little is known about the diversity of tropical animal communities in recently fireaffected environments. Here we assessed species richness, evenness, and community similarity of butterflies and odonates in landscapes located in unburned isolates and burned areas in a habitat mosaic that was severely affected by the 1997/98 ENSO (El Niñ o Southern Oscillation) event in east Kalimantan, Indonesian Borneo. In addition related community similarity to variation in geographic distance between sampling sites and the habitat/vegetation structure Species richness and evenness differed significantly among landscapes but there was no congruence between both taxa. The species richness of butterflies was, for example, highest in sites located in a very large unburned isolate whereas odonate species richness was highest in sites located in a small unburned isolate and once-burned forest. We also found substantial variation in the habitat/vegetation structure among landscapes but this was mainly due to variation between unburned and burned landscapes and variation among burned landscapes.Both distance and environment (habitat/vegetation) contributed substantially to explaining variation in the community similarity (beta diversity) of both taxa. The contribution of the environment was, however, mainly due to variation between unburned and burned landscapes, which contained very different assemblages of both taxa. Sites located in the burned forest contained assemblages that were intermediate between assemblages from sites in unburned forest and sites from a highly degraded slash-and-burn area indicating that the burned forest was probably recolonised by species from these disparate environments. We, furthermore, note that in contrast to species richness (alpha diversity) the patterns of community similarity (beta diversity) were highly congruent between both taxa. These results indicate that community-wide multivariate measures of beta diversity are more consistent among taxa and more reliable indicators of disturbance, such as ENSO-induced burning, than univariate measures.

Fossil butterflies are extremely rare. Yet, they are the only direct evidence of the first appearance of particular characters and as such, they are crucial for calibrating a molecular clock, from which divergence ages are estimated. In turn, these estimates, in combination with paleogeographic information, are most important in paleobiogeographic considerations. The key issue here is the correct allocation of fossils on the phylogenetic tree from which the molecular clock is calibrated.The allocation of a fossil on a tree should be based on an apomorphic character found in a tree based on extant species, similar to the allocation of a new extant species. In practice, the latter is not done, at least not explicitly, on the basis of apomorphy, but rather on overall similarity or on a phylogenetic analysis, which is not possible for most butterfly fossils since they usually are very fragmentary. Characters most often preserved are in the venation of the wings. Therefore, special attention is given to possible apomorphies in venational characters in extant butterflies. For estimation of divergence times, not only the correct allocation of the fossil on the tree is important, but also the tree itself influences the outcome as well as the correct determination of the age of the fossil. These three aspects are discussed.        All known butterfly fossils, consisting of 49 taxa, are critically reviewed and their relationship to extant taxa is discussed as an aid for correctly calibrating a molecular clock for papilionoid Lepidoptera. In this context some aspects of age estimation and biogeographic conclusions are briefly mentioned in review. Specific information has been summarized in four appendices.

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ISSN (online): 1802-8829http://www.eje.cz 6th, 2014. I feel it is appropriate to name the fossil after him. MATERIAL AND METHODSThe fossil ( Fig. 1) was found in the Fur formation on the island of Fur in the Limfjord in N Denmark. It is a 60 m thick marine deposit of diatoms and clay minerals with many layers of volcanic ash. It was formed shortly after the Paleocene to Eocene transition, about 55 Ma (Pedersen et al., 2004) and previously estimated at 57 Ma (Rust, 1998). The deposit is famous for its many fossils, not only marine animals but also terrestrial animals, which apparently were blown into the sea or actively fl ew over the water. Among the numerous insect fossils only the present fossil can be identifi ed as a butterfl y.The fossil is of a relatively broad-winged insect with wings folded down enclosing the legs. Outline of hindwings diffi cult to make out and most of the head is missing, but the dark lines in front of it may be remnants of palpi. Apex and greater part of termen of forewings are absent. Because of its fi ne detail the specimen in Fig. 1a is thought to be the compressed insect (although little is left except coloration of the substrate), of which the imprint is preserved in the counterpart illustrated in Fig. 1b. Therefore, the uppermost forewing is supposed to be the right wing. In the rest of the fossil the veins of four wings are lying on top of each other, making it diffi cult to decide which vein belongs to which wing. Part of the underlying (left) forewing is free and, although in this wing the apex is missing, several veins are clearly visible and there are no structures lying under it and blurring the picture. The venation of the "free" part of the (supposed) left forewing (right part of the fossil in Fig. 1a) was drawn on a transparent sheet and slid over the other forewing. By moving the Reconstructing a 55-million-year-old butterfl y (Lepidoptera: Hesperiidae) RIENK DE JONGNaturalis Biodiversity Center, PO Box 9517, 2300 RA Leiden, The Netherlands; e-mail: rienk.dejong@naturalis.nl Key words. Fossil, oldest butterfl y, Hesperiidae, Protocoeliades kristenseni, new genus, new species, divergence time, evolution, biogeography Abstract. The oldest butterfl y fossil known, which was formed about 55 Ma in what is now Denmark, is described. The fragments of its forewing venation indicates it belongs to the Hesperiidae. Further reconstruction indicates that it fi ts in the Coeliadinae and is close to the extant genera Hasora and Burara. It is here described as Protocoeliades kristenseni gen. et sp. n. It is the fi rst butterfl y fossil found on a continent (Europe) where its closest relatives do not currently occur. Its position on the phylogenetic tree of the Coeliadinae and its importance in understanding the time dimension in the evolution of butterfl ies, and their ecological and biogeographic implications are discussed. ZooBank Article LSID: E0B5F7AD-5352-493B-9016-F16A9C596D70

The Oriental butterfly genus Arhopala is by far the most species-rich genus within the subfamily Theclinae. We investigated whether biotic interactions with larval host plants or ants are phylogenetically constrained in the evolutionary history of Arhopala , by collating available information on the ecology of Arhopala from the literature as well as from personal observations, and analysing and interpreting these data rigorously in a phylogenetic context. Larvae of all species in the sister-group of Arhopala and Flos feed on Fabaceae. However, the predominant feeding preference of Arhopala s.l. , with the exception of a particular monophyletic and species-rich group, called 'Group A', appears to be centred on Fagaceae, with additional records of Dipterocarpaceae feeding. The preference for Fagaceae has strong phylogenetic inertia, as indicated by T-PTP tests designed to test for phylogenetic signal. Adding all available life-history data, using the phylogeny presented before as scaffolding, further increased the phylogenetic signal in host plant data. Feeding on Fabaceae (mainly in the outgroup) and Euphorbiaceae also showed significant phylogenetic signal, but feeding on Dipterocarpaceae did not and was found scattered across the phylogeny. Except for the Dipterocarpaceae, phylogenetic signal in feeding preferences was very clear, even despite uneven taxon sampling and apparent lability in host plant use. Ant association also demonstrated historical constraint, but based on the phylogenetic hypothesis presented here it was not clear whether increased ant association intimacy emerged independently various times in Group A, as well as in several basal groups. Our finding of distinct phylogenetic patterns in the host plant and to a lesser degree in ant association data contradicts the hypothesis that life-history traits are very labile in the Lycaenidae.