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FIGURE 4. A in Phylogeography, genetic structure and wing pattern variation of Erebia pronoe (Esper, 1780) (Lepidoptera: Nymphalidae) in Europe

FIGURE 4. A Maximum Likelihood (ML) tree of Erebia pronoe based on mitochondrial data of COI genes. Bootstrap support values are shown above the branches. A and B phylogroups are indicated on the right. The ML pattern is congruent with the statistical parsimony pattern of the sampled individuals. Sequences comprising E. pronoe populations from isolated clades are shown in grey shading. COI sequences of E. euryale (GU669656) and E. pandrose (KP870341) were used as outgroups. (a) The ML tree of E. pronoe based on mitochondrial data of CR markers. Bootstrap support values are shown above the branches. The ML pattern of CR markers is congruent with the ML pattern of COI genes.

FIGURE 5 in Phylogeography, genetic structure and wing pattern variation of Erebia pronoe (Esper, 1780) (Lepidoptera: Nymphalidae) in Europe

FIGURE 5. Scatterplot of canonical variate analysis. Morphometric overlap in Erebia pronoe forewings structure according to canonical variates CV1 and CV2. IT: Italy; SL: Slovenia; SP: Spain; SR: Slovakia.

FIGURE 3 in Phylogeography, genetic structure and wing pattern variation of Erebia pronoe (Esper, 1780) (Lepidoptera: Nymphalidae) in Europe

FIGURE 3. Statistical parsimony network of Erebia pronoe based on the COI mitochondrial dataset and statistical parsimony as implemented in TCS v1.21. Each haplotype is labelled by a number. Unsampled and extinct haplotypes are indicated by small circles. Division of the E. pronoe populations into seven haplogroups was detected, described in the Results section. These groups are illustrated with different symbols. Collection sites are indicated in the legend.

FIGURE 1 in Phylogeography, genetic structure and wing pattern variation of Erebia pronoe (Esper, 1780) (Lepidoptera: Nymphalidae) in Europe

FIGURE 1. Map of collection sites of Erebia pronoe in European mountain ranges and possible re-colonization paths of E. pronoe after the retreat of glacial stages during the Pleistocene. Grey shading and symbols represent different collection sites (by states); dashed arrows illustrate hypothesized paths of re-colonization from the refugia in the Pyrenees and Alps/Central Europe; triangles illustrate specimens collected from the European mountain ranges; circles illustrate specimens mined from the GenBank and BOLD databases. The number of analysed specimens is given in Table 1.

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Data from: Testing classical species properties with contemporary data: how 'bad species' in the brassy ringlets (Erebia tyndarus complex, Lepidoptera) turned good

All species concepts are rooted in reproductive, and ultimately genealogical, relations. Genetic data are thus the most important source of information for species delimitation. Current ease of access to genomic data and recent computational advances are blooming a plethora of coalescent-based species delimitation methods. Despite their utility as objective approaches to identify species boundaries, coalescent-based methods i) rely on simplified demographic models that may fail to capture some attributes of biological species, ii) do not make explicit use of the geographic information contained in the data, and iii) are often computationally intensive. In this paper we present a case of species delimitation in the Erebia tyndarus species complex, a taxon regarded as a classic example of problematic taxonomic resolution. Our approach to species delimitation used genomic data to test predictions rooted in the biological species concept and in the criterion of coexistence in sympatry. We 1) obtained RAD sequencing data from a carefully designed sample, 2) applied two genotype clustering algorithms to identify genetic clusters and 3) performed within-clusters and between-clusters analyses of isolation by distance (IBD) as a test for intrinsic reproductive barriers. Comparison of our results with those from a Bayes Factor Delimitation (BFD*) coalescent-based analysis, showed that coalescent-based approaches may lead to overconfident splitting of allopatric populations, and indicated that incorrect species delimitation is likely to be inferred when an incomplete geographic sample is analysed. While we acknowledge the theoretical justification and practical usefulness of coalescent-based species delimitation methods, our results stress that, even in the phylogenomic era, the toolkit for species delimitation should not dismiss more traditional, biologically grounded, approaches coupling genomic data with geographic information.

FIGURE 19 in A checklist of the satyrine genus Erebia (Lepidoptera) (1758–2006)

FIGURE 19: Another, rather more crudely painted, copy of Reiner and von Hohenwarth, published in Ulm, in 1793. This copy is held in the library of Stockholm University (photo: Clas-Ove Strandberg).

FIGURE 15 in A checklist of the satyrine genus Erebia (Lepidoptera) (1758–2006)

FIGURE 15: Dr Kenelm Philip, University of Fairbanks, Alaska, with some of the Smithsonian collection of nearctic Erebia butterflies. Knowledge of arctic butterflies has been much enhanced through the efforts of the Canadian Northern Insect Survey (1947–1957) and the Alaska Lepidoptera Survey (ALS), an ongoing project based at the University of Alaska (photo: John Tennent)

FIGURE 10 in A checklist of the satyrine genus Erebia (Lepidoptera) (1758–2006)

FIGURE 10: Warren and his wife at Urbachtal, Innertkirchen, Switzerland, on the 20th of August 1960 (photo: Elizabeth Warren).

FIGURE 4 in A checklist of the satyrine genus Erebia (Lepidoptera) (1758–2006)

FIGURE 4: The species recognised by most North American authors as Erebia theano at North Fork Pass, a famous locality near the bottom of the Dempster Highway, Canada, Yukon Territory (photo: John Tennent)

FIGURE 2 in A checklist of the satyrine genus Erebia (Lepidoptera) (1758–2006)

FIGURE 2: Erebia discoidalis—one of the most northerly nearctic species, photographed at Inuvik in Canada's Northwest Territory (photo: John Tennent)

FIGURE 14 in A checklist of the satyrine genus Erebia (Lepidoptera) (1758–2006)

FIGURE 14: Erebia ligea in the web of what is probably an orb-weaving spider (Araneidae) near Severobaikalsk, on the northwest coast of Lake Baikal, Siberia. The insect on the web is an ant of the Formica rufa group (Formicidae), which has presumably taken the opportunity of a potential free meal (photo: John Tennent)

FIGURE 11 in A checklist of the satyrine genus Erebia (Lepidoptera) (1758–2006)

FIGURE 11: Part of the Warren collection of Erebia butterflies in the BMNH, London (photo: John Tennent)

FIGURE 7 in A checklist of the satyrine genus Erebia (Lepidoptera) (1758–2006)

FIGURE 7: Erebia kefersteini between Mondy and Orlik in the East Sayan mountains, Siberia (photo: John Tennent)

FIGURE 9 in A checklist of the satyrine genus Erebia (Lepidoptera) (1758–2006)

FIGURE 9: Warren collecting butterflies at Val Tisch, Bergün, Switzerland, on the 29th of July 1952 (photo: Elizabeth Warren).

FIGURE 13 in A checklist of the satyrine genus Erebia (Lepidoptera) (1758–2006)

FIGURE 13: A crab spider (Thomisidae) of the genus Xysticus, with Erebia maurisius between Mondy and Orlik in the East Sayan mountains, Siberia (photo: John Tennent)

FIGURE 1 in A checklist of the satyrine genus Erebia (Lepidoptera) (1758–2006)

FIGURE 1: Erebia ligea photographed in the hills to the south west of Lake Baikal, Siberia. The parasitic red mite larvae attached to the butterfly's thorax belong to a species of Erythraeoidea or Trombidioidea. The precise impact on the host is not well studied, but is not known to result in host mortality. When fully fed, larvae drop to the ground to complete their life cycle. They are not infrequent on other satyrine butterflies, although in the author's experience it is unusual for them to be so prominent; they are more often attached to the lateral part of the thorax, below the wings. In one meadow near the shores of Lake Baikal infested specimens of E. ligea could be identified in flight even from a distance (photo: John Tennent)

FIGURE 18 in A checklist of the satyrine genus Erebia (Lepidoptera) (1758–2006)

FIGURE 18: The handcoloured folded plate depicting Erebia cassioides and other insects from Reiner and von Hohenwarth, published in Klagenfurt, in 1792. (photo: Photographic Unit, BMNH, London).

FIGURE 16 in A checklist of the satyrine genus Erebia (Lepidoptera) (1758–2006)

FIGURE 16: Dr Alexandr Lvovsky, Zoological Institute of St Petersburg, Russia, with some of the Erebia butterflies in the Institute, which includes type material from well-known early entomologists including Grum-Grshimaïlo, Eversmann and Alpheraky, in addition to type material described by contemporary workers (photo: John Tennent).

FIGURE 17 in A checklist of the satyrine genus Erebia (Lepidoptera) (1758–2006)

FIGURE 17: A selection of Erebia type specimens, mainly from the collection of Hans Fruhstorfer, in the BMNH, London (photo: John Tennent)

FIGURE 3 in A checklist of the satyrine genus Erebia (Lepidoptera) (1758–2006)

FIGURE 3: Erebia youngi photographed at Eagle Summit in Alaska, USA. Many butterflies in northerly latitudes spend long periods basking on bare ground or on rocks (photo: John Tennent)