Corresponding author: Pilar Jurado-Angulo (
Academic editor: Susanne Randolf
The Pyrenees are inhabited by scattered populations of earwigs of the genus
One of the most characteristic genera of
The species of C
The known distribution range of
With this main aim, the specific objectives of this work are: (i) to confirm the taxonomic identification of
Sampling was conducted in different localities of the Catalonian Pyrenees (Girona, Lleida), Andorra and Italy (Valle d’Aosta). A total of 104 specimens, 95 specimens of
Geographic distribution of
Dry-mounted specimens were examined under a stereomicroscopy. Live specimens were photographed in the field with a Nikon digital camera. Extended depth-of-focus images of dry-mounted specimens, were taken with a digital camera Nikon and a lens Nikon AF-S VR Micro-Nikkor 105mm f/2.8G IF-ED and the software Helicon Remote v. 3. 9. 11 and Helicon Focus v. 7. 6. 4. Male genitalia were extracted and studied following the protocol described in
Total DNA was obtained from six specimens (Table
Specimens used for DNA analyses with their corresponding MNCN Entomology Collection codes (or original publication) and GenBank accession numbers.
Species | Specimen code | Geographic origin | Coordinates | GenBank COI |
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MNCN_Ent 296011 | Italy: Valle d’Aosta: Val Veny: Pré de Pascal |
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MNCN_Ent 296012 | Italy: Valle d’Aosta: Val Veny: Pré de Pascal |
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MNCN_Ent 296013 | Spain: Girona: La Molina: Tossa d’Alp |
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MNCN_Ent 296014 | Spain: Girona: La Molina: Tossa d’Alp |
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MNCN_Ent 296015 | Andorra: Sant Julià de Lòria: Bixessarri: Coll de la Gallina |
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MNCN_Ent 296016 | Andorra: Sant Julià de Lòria: La Rabassa |
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France: Chalmazel ( |
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5b-1 | Switzerland, Valais, Col du Grand Saint-Bernard, Liddes, 2160 m |
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5b-5 | Switzerland, Valais, Col du Grand Saint-Bernard, Liddes, 2160 m |
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2a-1 | Italy, Piedmont (Biella), Pennine Alps, Lago del Mucrone, Oropa, 1910 m |
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2a-3 | Italy, Piedmont (Biella), Pennine Alps, Lago del Mucrone, Oropa, 1910 m |
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17-1 | Italy, Lombardy (Sondrio), Western Rhaetian Alps, Franscia, Lanzada, 1480 m |
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18 | Italy, Lombardy (Sondrio), Western Rhaetian Alps, Franscia, Lanzada, 1480 m |
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Italy: Trento ( |
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Slovakia: Poľana ( |
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Portugal: Monchique ( |
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Mongolia: Ikh-tamir ( |
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The
The obtained
Evolutionary (taxonomic) units within
Based on the phylogenetic analyses, studied specimens of
Uncorrected (p) pairwise genetic distance matrix between specimens (short – long cerci) and taxa used in the phylogenetic analyses.
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Valle d’Aosta | Valais | Biella | Sondrio | Long cerci | Short cerci | ||||||||
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Valle d’Aosta | 0.00 | ||||||||||
Valais | 0.00-0.01 | 0.00 | |||||||||||
Biella | 0.08-0.09 | 0.08-0.09 | 0.02 | ||||||||||
Sondrio | 0.09 | 0.08-0.09 | 0.03-0.04 | 0.00 | |||||||||
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Long cerci | 0.19-0.21 | |||||||||||
Short cerci | 0.19-0.22 | 0.00-0.03 | 0.02-0.03 | ||||||||||
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0.18-0.21 | 0.15 | 0.14-0.15 | ||||||||||
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0.20-0.21 | 0.21 | 0.20-0.21 | 0.20 | ||||||||
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0.20-0.21 | 0.19 | 0.19-0.20 | 0.21 | 0.15 | ||||||||
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0.22-0.23 | 0.22 | 0.22 | 0.22 | 0.20 | 0.20 | ||||||
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0.24-0.25 | 0.20 | 0.20 | 0.21 | 0.23 | 0.22 | 0.19 |
Bayesian phylogenetic tree based on
Pyrenean specimens (Andorra and Girona) form a monophyletic group of poorly differentiated sequences (uncorrected p distance ranging from 0 to 0.03) (Fig.
Male specimens included in the Pyrenean clade (Girona and Andorra) present large variability in the shape of the cerci. Cerci range from long, almost straight convergent cerci (Figs
Cerci variation in
The specimens studied from the Alpine clade present long cerci with little curvature, cylindrical apically, progressively narrowed towards the apex and the inner margins without teeth or with one tooth. The diameter of the cerci of those specimens studied is generally smaller than that of the specimens of the Pyrenean clade. However, our sample is not representative of the variability already reported for the Alpine clade (
Live specimens of
The Pyrenean clade is sister to the single sequence representing the Massif Central clade (PP = 0.87). The genetic distance between Pyrenean and Massif Central populations is quite large (uncorrected p distance = 0.14–0.15). Among the large series of Pyrenean specimens studied we did not find the cerci morphology described for
Male genitalia from specimens of the Pyrenees, Alpine and Massif Central clades, including the lectotype of
Live specimens of
There is a strict correspondence between mtDNA clades and geographic areas, with all samples from the Pyrenees included in a well-supported clade, sister to the Massif Central specimen, and those, in turn, related to the Alpine specimens. Sequences of the specimens from Tossa d’Alp (with short and long cerci respectively) are closer to each other than to the short cerci specimens from Andorra, therefore, at the molecular level, specimens with short and long cerci from the Pyrenees correspond to a single taxon. Results from the nymphs raised under controlled conditions, with adult males including long (see for example MNCN_Ent 296013; Fig.
Populations of
Intraspecific variability of morphological structures, as pygidium or cerci, is well known in earwigs (
The taxonomic implications of the large shape variability in male cerci need to be addressed in the case of Pyrenean
Most of the confusion derived from the early synonymization of
We thank Pilar Pavón Gozalo, Vladimir Salvador de Jesús and Gonzalo García for their help during field surveys. This work was possible thanks to Mercedes París, curator of Entomology of the Museo Nacional de Ciencias Naturales (Madrid), who helped us during the revision of the dry-preserved materials from the Entomological Collection and for assistance while taking images. We thank Paolo Fontana, Jose Luis Ruiz, and an additional anonymous reviewer for relevant suggestions that improved this manuscript. We also thank Markéta Kirstová and Petr Kočárek for comments and suggestions on this project. Thanks to Alberto Sánchez Vialas for help with the distribution map and to Fernando García Guerrero for help with the photographs of the male genitalia. This work was partially funded by the project grant PID2019-110243GB-100 / AEI/