Data Paper |
Corresponding author: Martin Wiemers ( martin.wiemers@senckenberg.de ) Academic editor: Rodolphe Rougerie
© 2020 Martin Wiemers, Nicolas Chazot, Christopher W. Wheat, Oliver Schweiger, Niklas Wahlberg.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Wiemers M, Chazot N, Wheat CW, Schweiger O, Wahlberg N (2020) A complete time-calibrated multi-gene phylogeny of the European butterflies. ZooKeys 938: 97-124. https://doi.org/10.3897/zookeys.938.50878
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With the aim of supporting ecological analyses in butterflies, the third most species-rich superfamily of Lepidoptera, this paper presents the first time-calibrated phylogeny of all 496 extant butterfly species in Europe, including 18 very localised endemics for which no public DNA sequences had been available previously. It is based on a concatenated alignment of the mitochondrial gene COI and up to eleven nuclear gene fragments, using Bayesian inferences of phylogeny. To avoid analytical biases that could result from our region-focussed sampling, our European tree was grafted upon a global genus-level backbone butterfly phylogeny for analyses. In addition to a consensus tree, the posterior distribution of trees and the fully concatenated alignment are provided for future analyses. Altogether a complete phylogenetic framework of European butterflies for use by the ecological and evolutionary communities is presented.
Butterflies of Europe, divergence times, macroecology, phylogeny, time tree
The incorporation of phylogenetic information in ecological theory and research has led to significant advancements by facilitating the connection of large-scale and long-term macro-evolutionary processes with ecological processes in the analysis of species interactions with their abiotic and biotic environments (
Although the amount of molecular data has increased exponentially during the last decades, most available phylogenetic studies are either restricted to a selected subset of species, higher taxa, or to small geographic areas. Complete and dated species-level phylogenetic hypotheses for species-rich taxa of larger regions have been restricted to vascular plants (
Here, we present the first comprehensive time-calibrated molecular phylogeny of all 496 extant European butterfly species (Lepidoptera: Papilionoidea), based on one mitochondrial and up to eleven nuclear genes, and the most recent systematic list of European butterflies (
Compared to other groups of insects, the phylogenetic relationships of butterflies are reasonably well-known, with robust backbone molecular phylogenies at the subfamily (
We analyse a dataset comprising all extant European species of butterflies (Papilionoidea), including the families Papilionidae, Hesperiidae, Pieridae, Lycaenidae, Riodinidae, and Nymphalidae. We base our species concepts, as well as the area defined as Europe, on the latest checklist of European butterflies (
The data were mainly collated from published sources and downloaded from NCBI GenBank (Suppl. material
In many cases, new sequences were generated for this study. For these specimens, protocols followed
Almost all genera are represented by multiple genes, except Borbo, Gegenes, Laeosopis, Callophrys, and Cyclyrius (the latter recently synonymised with Leptotes;
Newly sequenced species for which no published sequences had previously been available.
Taxon | Origin | COI | EF-1α | GAPDH | Wingless |
---|---|---|---|---|---|
Coenonympha orientalis | Greece | MN829478 | MN829462 | ||
Glaucopsyche paphos | Cyprus | MN829481 | MN829463 | ||
Gonepteryx maderensis | Portugal: Madeira | MN829482 | MN829464 | ||
Hipparchia azorina | Portugal: Azores | MN829483 | MN829465 | ||
Hipparchia bacchus | Spain: Canary Islands | MN829484 | MN829466 | ||
Hipparchia cretica | Greece: Crete | MN752718 | MN829467 | MN752786 | MN752837 |
Hipparchia gomera | Spain: Canary Islands | MN829485 | MN829468 | ||
Hipparchia maderensis | Portugal: Madeira | MN829486 | |||
Hipparchia mersina | Greece: Samos | MN752720 | MN829469 | MN752785 | MN752836 |
Hipparchia miguelensis | Portugal: Madeira | MN829487 | |||
Hipparchia sbordonii | Italy: Pontine Islands | MN752723 | |||
Hipparchia tamadabae | Spain: Canary Islands | MN829488 | |||
Hipparchia tilosi | Spain: Canary Islands | MN829489 | |||
Hipparchia wyssii | Spain: Canary Islands | MN829490 | MN829470 | ||
Lycaena bleusei | Spain | MN829492 | |||
Pieris balcana | North Macedonia | KC462788 | |||
Pieris wollastoni | Portugal: Madeira | KC462820 | |||
Thymelicus christi | Spain: Canary Islands | MN829496 |
A biogeographically restricted tree of a given taxon is inherently very asymmetrically sampled. To avoid potentially strong biases when estimating topology and divergence times we chose to build upon the recent genus-level tree of butterflies (
To estimate a time-calibrated tree of European butterflies, we first identified the position of the European lineages and designed a grafting procedure accordingly. We split the European butterflies that needed to be added to the tree into 12 subclades. For each of these subclades we combined the DNA sequences of the taxa already included in the backbone to the DNA sequences of the European taxa to assemble an aligned molecular matrix. After identifying the best partitioning scheme, we performed a tree reconstruction without time-calibration (i.e., only estimating branch lengths proportional to relative time). The subclade trees were then rescaled using the ages estimated in the backbone and were subsequently grafted. This procedure was repeated using 1000 trees from BEAST posterior distributions of the backbone and subclade trees in order to obtain a posterior distribution of grafted trees. The details of these procedures are described below.
The time-calibrated backbone tree provided by
The subclades, sorted by families, were defined as follows:
Papilionidae – All Papilionidae were placed into one subclade.
Hesperiidae – We identified two main clades to graft within the Hesperiidae: Hesperiinae and Pyrginae. The Hesperiinae subclade was extended to also encompass the subfamilies Heteropterinae and Trapezitinae. The genus Muschampia, not available in the backbone, was included in the Pyrginae subclade.
Pieridae – All Pieridae were considered as a single clade.
Lycaenidae – All Lycaenidae were considered as a single clade.
Riodinidae – The only European Riodinidae species, Hamearis lucina, was already available in the backbone tree.
Nymphalidae – European Nymphalidae were divided into seven subclades. (i) A subclade for the Apaturinae. (ii) In order to add Danaus chrysippus we generated a tree of Danainae. (iii) We combined the sister clades Heliconiinae and Limenitidinae into a single subclade. (iv) Nymphalinae was treated as a single subclade. (v) A first clade of Satyrinae contained the genera Kirinia, Pararge, Lasiommata, Tatinga, Chonala and Lopinga. (vi) A second Satyrinae clade contained the genera Calisto, Euptychia, Callerebia, Proterebia, Gyrocheilus, Strabena, Ypthima, Ypthimomorpha, Stygionympha, Cassionympha, Neocoenyra, Pseudonympha, Erebia, Boerebia, Hyponephele, Cercyonis, Maniola, Aphantopus, Pyronia, Faunula, Grumia, Paralasa, Melanargia, Hipparchia, Berberia, Oeneis, Neominois, Karanasa, Brintesia, Arethusana, Satyrus, Pseudochazara, and Chazara. (vii) A third Satyrinae clade was created for the genus Coenonympha. Charaxinae were not treated separately from the backbone. Charaxes jasius is the only Charaxinae occurring in Europe and Charaxes castor (which is very closely related to C. jasius;
For each subclade we ran PartitionFinder 2.1.1 (
For each subclade, the dataset was imported in BEAUTi 1.8.3 (
Subclades were grafted on the backbone as follows. One backbone was sampled from the posterior distribution of time-calibrated trees from
We describe below the details of the phylogenetic tree reconstruction for each subclade.
1. Papilionidae
Dataset – The dataset for the Papilionidae consisted of 36 taxa to which three outgroups were added: Macrosoma tipulata (Hedylidae), Achlyodes busiris (Hesperiidae), Pieris rapae (Pieridae). We concatenated 11 gene fragments (COI, CAD, EF-1α, GAPDH, ArgK, IDH, MDH, RpS2, RpS5, DDC, wingless).
PartitionFinder – PartitionFinder identified 12 subsets (Suppl. material
BEAST analysis – In order to improve the quality of our runs we replaced the default priors for rates of substitutions by uniform prior ranging between 0 and 10 for the following cases: subset5.at, subset5.cg, subset7.cg, subset7.gt, subset12.cg, subset12.gt. We used one molecular clock per subset identified by PartitionFinder and obtained good mixing and convergence. We used a birth-death tree prior. We performed three runs of 40 million generations, sampling trees and parameters every 4000 generations.
Grafting – For grafting, the outgroups were removed, as well as Baronia brevicornis, the first Papilionidae to diverge and endemic to Mexico (
2. Hesperiidae: Hesperiinae
Dataset – The dataset for the Hesperiinae consisted of 169 taxa to which two outgroups were added: Typhedanus ampyx (Hesperiidae: Eudaminae), Mylon pelopidas (Hesperiidae: Pyrginae). We concatenated 10 gene fragments (COI, CAD, EF-1α, GAPDH, ArgK, IDH, MDH, RpS2, RpS5, wingless).
PartitionFinder – PartitionFinder identified 17 subsets (Suppl. material
BEAST analysis – Preliminary analyses showed problems with the subset 3 (ArgKin_pos3) which was therefore removed from the analyses. In order to improve the quality of our runs we replaced the default priors for rates of substitutions by uniform priors ranging between 0 and 10 for the following case: subset17.cg. The substitution model for the subset 14 was also changed into HKY+I after preliminary analyses. We used one molecular clock per subset identified by PartitionFinder and obtained good mixing and convergence. We used a birth-death tree prior. We performed two runs of 150 million generations, sampling trees and parameters every 15000 generations.
Grafting – For grafting, the outgroups were removed and the subclade grafted at the MRCA of Hesperiinae.
3. Hesperiidae: Pyrginae
Dataset – The dataset for the Pyrginae consisted of 77 taxa to which three outgroups were added: Typhedanus ampyx (Hesperiidae: Eudaminae), Pyrrhopyge zenodorus (Hesperiidae: Pyrginae), and Hasora khoda (Hesperiidae: Coeliadinae). We concatenated ten gene fragments (COI, CAD, EF-1α, GAPDH, ArgK, IDH, MDH, RpS2, RpS5, wingless).
PartitionFinder – PartitionFinder identified 14 subsets (Suppl. material
BEAST analysis – In order to improve the quality of our runs we replaced the default priors for rates of substitutions by uniform priors ranging between 0 and 10 for the following cases: subset7.ac, subset7.gt, subset14.cg, subset3.cg. Preliminary analyses showed problems when using a separate molecular clock for each subset identified by PartitionFinder. We restricted the analysis to one molecular clock. We used a birth-death tree prior. We performed two runs of 100 million generations, sampling trees and parameters every 10000 generations.
Grafting – For grafting, the outgroups were removed, and the subclade grafted at the MRCA of Pyrginae.
4. Pieridae
Dataset – The dataset for the Pieridae consisted of 126 taxa to which three outgroups were added: Bicyclus anynana (Nymphalidae), Achlyodes busiris (Hesperiidae), and Papilio glaucus (Papilionidae). We concatenated eleven gene fragments (COI, CAD, EF-1α, GAPDH, ArgK, IDH, MDH, RpS2, RpS5, DDC, wingless).
PartitionFinder – PartitionFinder identified 17 subsets (Suppl. material
BEAST analysis – In order to improve the quality of our runs we replaced the default priors for rates of substitutions by uniform priors ranging between 0 and 10 for the following case: subset7.cg. The substitution model for the subset 7 was also changed into GTR+G after preliminary analyses. We used one molecular clock per subset identified by PartitionFinder and obtained good mixing and convergence. We used a birth-death tree prior. We performed two runs of 100 million generations, sampling trees and parameters every 10000 generations.
Grafting – For grafting, the outgroups were removed, and the subclade grafted at the MRCA of Pieridae.
5. Lycaenidae
Dataset – The dataset for the Lycaenidae consisted of 187 taxa to which three outgroups were added: Bicyclus anynana (Nymphalidae), Pieris rapae (Pieridae) and Hamearis lucina (Riodinidae). We concatenated 12 gene fragments (COI, CAD, EF-1α, GAPDH, ArgK, IDH, MDH, RpS2, RpS5, DDC, wingless and H3).
PartitionFinder – PartitionFinder identified 12 subsets (Suppl. material
BEAST analysis – In order to improve the quality of our runs we replaced the default priors for rates of substitutions by uniform priors ranging between 0 and 10 for the following cases: subset3.cg, subset6.ag, subset6.at, subset11.gt_subst7.cg. We used one molecular clock per subset identified by PartitionFinder and obtained good mixing and convergence. We used a birth-death tree prior. We performed two runs of 150 million generations, sampling trees and parameters every 15000 generations.
Grafting – For grafting, the outgroups were removed, and the subclade grafted at the MRCA of Lycaenidae.
6. Nymphalidae: Danainae
Dataset – The dataset for the Danainae consisted of 7 taxa to which two outgroups were added: Euploea camaralzeman (Nymphalidae: Danainae) and Lycorea halia (Nymphalidae: Danainae). We concatenated 9 gene fragments (COI, CAD, EF-1α, GAPDH, IDH, MDH, RpS2, RpS5, wingless).
PartitionFinder – PartitionFinder identified eight subsets (Suppl. material
BEAST analysis – We used one molecular clock per subset identified by PartitionFinder and obtained good mixing and convergence. We used a birth-death tree prior. We performed two runs of 20 million generations, sampling trees and parameters every 2000 generations.
Grafting – For grafting, the outgroups were removed, and the subclade grafted at the MRCA of Danainae.
7. Nymphalidae: Apaturinae
Dataset – The dataset for the Apaturinae consisted of nine taxa to which two outgroups were added: Timelaea albescens (Nymphalidae: Apaturinae) and Biblis hyperia (Nymphalidae: Biblidinae). We concatenated ten gene fragments (COI, CAD, EF-1α, GAPDH, ArgK, IDH, MDH, RpS2, RpS5, wingless).
PartitionFinder – PartitionFinder identified seven subsets (Suppl. material
BEAST analysis – We used one molecular clock per subset identified by PartitionFinder and obtained good mixing and convergence. We used a birth-death tree prior. We performed two runs of 20 million generations, sampling trees and parameters every 2000 generations.
Grafting – For grafting, the outgroups were removed, and the subclade grafted at the MRCA of Danainae.
8. Nymphalidae: Heliconiinae + Limenitidinae
Dataset – The dataset combined the sister clades Heliconiinae and Limenitidinae and consisted of 92 taxa to which three outgroups were added: Amnosia decora (Nymphalidae: Pseudoergolinae), Apatura iris (Nymphalidae: Apaturinae) and Libythea celtis (Nymphalidae: Libytheinae). We concatenated eleven gene fragments (COI, CAD, EF-1α, GAPDH, ArgK, IDH, MDH, RpS2, RpS5, DDC, wingless).
PartitionFinder – PartitionFinder identified 14 subsets (Suppl. material
BEAST analysis – Preliminary analyses showed problems with the subset 14 (RpS2_pos2) which was therefore removed from the analyses. In order to improve the quality of our runs we replaced the default priors for rates of substitutions by uniform priors ranging between 0 and 10 for the following case: subset7.cg. We used one molecular clock per subset identified by PartitionFinder and obtained good mixing and convergence. We used a birth-death tree prior. We performed two runs of 100 million generations, sampling trees and parameters every 10000 generations.
Grafting – For grafting, the outgroups were removed, and the subclade grafted at the split between Limenitidinae and Heliconiinae.
9. Nymphalidae: Nymphalinae
Dataset – The dataset of Nymphalinae consisted of 83 taxa to which two outgroups were added: Historis odius (Nymphalidae: Nymphalinae) and Pycina zamba (Nymphalidae: Nymphalinae). We concatenated eleven gene fragments (COI, CAD, EF-1α, GAPDH, ArgK, IDH, MDH, RpS2, RpS5, DDC, wingless).
PartitionFinder – PartitionFinder identified 12 subsets (Suppl. material
BEAST analysis – In order to improve the quality of our runs we replaced the default priors for rates of substitutions by uniform priors ranging between 0 and 10 for the following case: subset5.cg. Preliminary analyses revealed problems when using one molecular clock per subset identified by Partition Finder. We restricted the analysis to one molecular clock for the mitochondrial gene fragments and one molecular clock for the nuclear gene fragments. We used a birth-death tree prior. We performed two runs of 100 million generations, sampling trees and parameters every 10000 generations.
Grafting – For grafting, the outgroups were removed, and the subclade grafted at the MRCA of Nymphalinae.
10. Nymphalidae: Satyrinae 1
Dataset – The first Satyrinae dataset consisted of 13 taxa, belonging to the genera Kirinia, Pararge, Lasiommata, Tatinga, Chonala, and Lopinga, to which three outgroups were added: Bicyclus anynana (Nymphalidae: Satyrinae), Acrophtalmia leuce (Nymphalidae: Satyrinae), and Ragadia makuta (Nymphalidae: Satyrinae). We concatenated 5 gene fragments (COI, EF-1α, GAPDH, RpS5, wingless).
PartitionFinder – PartitionFinder identified six subsets (Suppl. material
BEAST analysis – We used one molecular clock per subset identified by PartitionFinder and obtained good mixing and convergence. We used a birth-death tree prior. We performed two runs of 20 million generations, sampling trees and parameters every 2000 generations.
Grafting – For grafting, the outgroups were removed, and the subclade grafted at the crown of the clade after removing the outgroups.
11. Nymphalidae: Satyrinae 2
Dataset – The second Satyrinae dataset consisted of 161 taxa, belonging to the genera Calisto, Euptychia, Callerebia, Proterebia, Gyrocheilus, Strabena, Ypthima, Ypthimomorpha, Stygionympha, Cassionympha, Neocoenyra, Pseudonympha, Erebia, Boerebia, Hyponephele, Cercyonis, Maniola, Aphantopus, Pyronia, Faunula, Grumia, Paralasa, Melanargia, Hipparchia, Berberia, Oeneis, Neominois, Karanasa, Brintesia, Arethusana, Satyrus, Pseudochazara, and Chazara, to which three outgroups were added: Coenonympha pamphilus (Nymphalidae: Satyrinae), Taygetis virgilia (Nymphalidae: Satyrinae), and Pronophila thelebe (Nymphalidae: Satyrinae). We concatenated ten gene fragments (COI, CAD, EF-1α, GAPDH, ArgK, IDH, MDH, RpS2, RpS5, wingless).
PartitionFinder – PartitionFinder identified eleven subsets (Suppl. material
BEAST analysis – In order to improve the quality of our runs we replaced the default priors for rates of substitutions by uniform prior ranging between 0 and 10 for the following cases: subset5.ac, subset5.ag, subset5.at, subset5.cg, subset5.gt. We used one molecular clock per subset identified by PartitionFinder and obtained good mixing and convergence. We used a birth-death tree prior. We performed two runs of 100 million generations, sampling trees and parameters every 10000 generations.
Grafting – For grafting, the outgroups were removed, and the subclade grafted at the crown of the clade after removing the outgroups.
12. Nymphalidae: Satyrinae 3
Dataset – The third Satyrinae dataset consisted of 15 taxa all belonging to the genus Coenonympha, to which two outgroups were added: Sinonympha amoena (Nymphalidae: Satyrinae) and Oressinoma sorata (Nymphalidae: Satyrinae). We concatenated nine gene fragments (COI, CAD, EF-1α, GAPDH, IDH, MDH, RpS2, RpS5, wingless).
PartitionFinder – PartitionFinder identified six subsets (Suppl. material
BEAST analysis – We used one molecular clock per subset identified by PartitionFinder and obtained good mixing and convergence. We used a birth-death tree prior. We performed two runs of 20 million generations, sampling trees and parameters every 2000 generations.
Grafting – For grafting, the outgroups were removed, and the subclade grafted at the crown of Coenonympha.
Species identities of the chosen sequences for the dataset were validated by blasting the DNA barcode sequences against the Barcode Of Life Database (http://www.boldsystems.org/), which has a good representation of European butterfly species due to a number of barcoding projects implemented in different countries (e.g.,
We estimated our time-calibration from a recent re-evaluation of the timing of divergence of higher-level Papilionoidea. We used the topology inferred by
We show here a synthetic tree summarising the posterior distribution of topologies and node ages, but the posterior distribution of grafted trees can be found in the Supporting Information, providing a distribution of alternative topologies and node ages estimated by BEAST. We strongly advise any researcher using these phylogenetic trees to repeat any analyses on at least 100 trees randomly sampled from this posterior distribution in order to account for topology and node age uncertainties. This tree can also help to identify the sections of the tree lacking molecular information and therefore points at the sections that should be targeted in the future when generating new molecular data.
The analysed dataset (a concatenated alignment of the genes COI, CAD, EF-1α, GAPDH, ArgK, IDH, MDH, RpS2, RpS5, DDC, wingless, and H3) is available in NEXUS format and the posterior distribution of ML trees and the consensus tree in NEWICK format at DOI: https://dx.doi.org/10.5281/zenodo.3531555.
We have generated a robust phylogenetic hypothesis for all European species of butterflies with estimations of divergence times (Fig.
Proposal for changes in the current taxonomic checklist by
Current name ( |
Proposed name ( |
Muschampia cribrellum (Eversmann, 1841) | Favria cribrellum (Eversmann, 1841) |
Carcharodus lavatherae (Esper, 1783) | Muschampia (Reverdinus) lavatherae (Esper, 1783) |
Carcharodus orientalis Reverdin, 1913 | Muschampia (Reverdinus) orientalis (Reverdin, 1913) |
Carcharodus floccifera (Zeller, 1847) | Muschampia (Reverdinus) floccifera (Zeller, 1847) |
Carcharodus stauderi Reverdin, 1913 | Muschampia (Reverdinus) stauderi (Reverdin, 1913) |
Carcharodus baeticus (Rambur, 1839) | Muschampia (Reverdinus) baeticus (Rambur, 1840) |
We thank Fabien L. Condamine (CNRS Montpellier) for his comments on a previous version of this manuscript which helped to improve the paper. MW thanks Brigitte Gottsberger (University of Vienna) for assistance in the lab and the following colleagues for specimen samples or sequences: Benedicto Acosta-Fernandez (Spain), Bernard Turlin (France), Dirk Gerber (Germany), Eddie John (UK), John Coutsis (Greece), Javier García (Spain), Karen van Dorp (Netherlands), Klaus Schurian (Germany), Pedro Oromí (Spain), Peter Russell (UK), Roger Vila (Spain), Vlad Dinca (Finland), Xavier Merit (France), Zdenek Fric (Czech Republic), and Zdravko Kolev (Bulgaria). We thank Pascal Wiemers of Selam-X (http://www.selam-x.com/studio) for the help in producing Figure
Table listing of European butterfly species with higher taxonomy, voucher codes and accession numbers for the sequences used to build the phylogeny.
Data type: List of taxa with accession numbers
Table S1–S12
Data type: PartitionFinder results
Explanation note: Table S1. PartitionFinder results for dataset 1 (Papilionidae). Table S2. PartitionFinder results for dataset 2 (Hesperiidae: Hesperiinae). Table S3. PartitionFinder results for dataset 3 (Hesperiidae: Pyrginae). Table S4. PartitionFinder results for dataset 4 (Pieridae). Table S5. Partitionfinder results for dataset 5 (Lycaenidae). Table S6. PartitionFinder results for dataset 6 (Nymphalidae: Danainae). Table S7. PartitionFinder results for dataset 7 (Nymphalidae: Apaturinae). Table S8. PartitionFinder results for dataset 8 (Nymphalidae: Heliconiinae + Limenitidinae). Table S9. PartitionFinder results for dataset 9 (Nymphalidae: Nymphalinae). Table S10. PartitionFinder results for dataset 10 (Nymphalidae: Satyrinae 1). Table S11. PartitionFinder results for dataset 11 (Nymphalidae: Satyrinae 2). Table S12. PartitionFinder results for dataset 12 (Nymphalidae: Satyrinae 3).
Figure S1. Time-calibrated tree of European butterflies
Data type: Phylogenetic dendrogram
Explanation note: Unlabelled terminal branches (with monophyletic entities collapsed) represent non-European taxa which were included in the global backbone tree. Age estimates are indicated at the nodes (Ma). Node bars represent the 95% credibility intervals.
Figure S2. Time-calibrated tree of European butterflies Section I: Papilionidae, Hesperiidae & Pieridae
Data type: Phylogenetic dendrogram
Explanation note: Age estimates are indicated at the nodes (Ma).
Figure S3. Time-calibrated tree of European butterflies Section II. Riodinidae & Lycaenidae.
Data type: Phylogenetic dendrogram
Explanation note: Age estimates are indicated at the nodes (Ma).
Figure S4. Time-calibrated tree of European butterflies Section III: Nymphalidae Part I: Subfamilies Limenitidinae, Heliconiinae, Apaturinae & Nymphalinae
Data type: Phylogenetic dendrogram
Explanation note: Age estimates are indicated at the nodes (Ma).
Figure S5. Time-calibrated tree of European butterflies Section IV: Nymphalidae Part II: Subfamilies Libytheinae, Danainae & Satyrinae
Data type: Phylogenetic dendrogram
Explanation note: Age estimates are indicated at the nodes (Ma).
Figure S6. Time-calibrated tree of European butterflies
Data type: Phylogenetic dendrogram
Explanation note: Unlabelled terminal branches (with monophyletic entities collapsed) represent non-European taxa which were included in the global backbone tree. Posterior probabilities are indicated at the nodes (Ma).