Dtsch. Entomol. Z. 68 (2) 2021, 235-248 | DOI 10.3897/dez.68.68020 ee BERLIN The Pyrenean species of Chelidura (Dermaptera, Forficulidae) Pilar Jurado-Angulo', Yolanda Jiménez-Ruiz', Mario Garcia-Paris? 1 Department of Biodiversity and Evolutionary Biology. Museo Nacional de Ciencias Naturales, MNCN-CSIC. c/ José Gutiérrez Abascal, 2. 28006, Madrid. Spain http://zoobank.org/43AD7562-7AF5-426D-A7EF-17225DD9AD98 Corresponding author: Pilar Jurado-Angulo (si.pilarjurado@gmail.com) Academic editor: Susanne Randolf # Received 28 April 2021 Accepted 6 July 2021 Published 10 August 2021 Abstract The Pyrenees are inhabited by scattered populations of earwigs of the genus Chelidura Latreille, 1825. There is some controversy about the specific assignment of these populations: while most authors assign them to C. pyrenaica (Gené, 1832), other consider that C. aptera (Mégerlé, 1825) is also present in the Pyrenees. The main objective of this work was to revise the identity and synonyms of Pyrenean Chelidura. Specimens from recent fieldwork and collections (MNCN-CSIC) were used for morphological and molecular studies (cytochrome oxidase 1). All Pyrenean specimens shared similar cox/ sequences, very divergent from those of Alpine C. ap- tera. As a consequence, the variability observed in male cerci morphology from the Pyrenees, ranging from long and slightly curved to short and very curved, corresponded to C. pyrenaica, and the presence of C. apfera in the Pyrenees can be rejected. As previously suggested by Maccagno (1933) and Fontana et al. (2021), the revision of the synonymic list uncovered the misplacement of the name F: simplex Germar, 1825 under the synonymy of C. apfera, while it rather represents a synonym of C. pyrenaica (syn. nov.). Forficula simplex has nomenclatural priority over C. pyrenaica, however both names meet the requirements of the article 23.9.1 of the International Code of Zoological Nomenclature to retain the prevailing usage of C. pyrenaica (nomen protectum) over I. simplex (nomen oblitum). Additionally, we discuss the taxonomic status of Chelidura arverna David & Van Herrewege, 1973 stat. nov. from the French Massif Central. Key Words Chelidura aptera, Chelidura arverna, Chelidura pyrenaica, Cytochrome oxidase 1, earwigs, geographic distribution, intraspecific variation, morphology, taxonomy Introduction One of the most characteristic genera of Dermaptera in the high elevations of the European Mountains is Che- lidura Latreille, 1825, represented by robust large-sized species often found in the upper limit of the coniferous forests. After the recent revision by Kirstova et al. (2020) who reconsidered the status of Mesochelidura Verhoeff, 1902 and Chelidurella Verhoeff, 1902, previously syn- onymized with Chelidura by Steinmann (1993), the ge- nus Chelidura includes 13 species confined to mountains of the Palaearctic region (Koéarek 2004; Kirstova et al. 2020). However, the genus is still in need of a rigorous taxonomic revision to determine the status of the Asian species (Kirstova et al. 2020). The species of Chelidura are characterized by absence of wings, a broad and large body with rudimentary tegmi- na, abdomen strongly dilated towards the posterior end, and flat, rounded and not protruding pygidium (Albouy and Caussanel 1990; Koéarek 2004; Kirstova et al. 2020; Fontana et al. 2021). The Pyrenean Mountain Chain is inhabited by scattered populations of Chelidura, distrib- uted over the Spanish, Andorran and French sides of the chain (Lapeira and Pascual 1980; Albouy and Caussanel 1990; Herrera-Mesa 1999; Fontana et al. 2021). The spe- cific ascription of these Pyrenean populations is subject Copyright Pilar Jurado-Angulo et al. 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. 236 to discussion. Some authors only mentioned the presence of C. pyrenaica (Gené, 1832) in the Pyrenees (Marquet 1877; Finot 1890; de Bormans and Krauss 1900; Houl- bert 1900; Xambeu 1903; Kirby 1904; Burr 1904; Xam- beu 1907; Hamon 1956; Popham 1968; Vancassel and Foraste 1980; Dauphin 1987; Fontana 1999; Dusoulier 2004; Fontana et al. 2021), while some others considered that C. pyrenaica and also C. aptera (Megerle, 1825), are both present in the Pyrenees (Serville 1839; Fieber 1853; Bolivar 1878; Cazurro Ruiz 1888; Chopard 1922; Chopard 1951; Boeseman 1954; Amiet 1961; Sakai 1973; Harz and Kaltenbach 1976; Lapeira and Pascual 1980; Caussanel and Albouy 1987; Albouy and Caussanel 1990; Herrera-Mesa 1999). Chelidura pyrenaica and C. aptera are two Europe- an species with ecological similarities. Both species are found in mountains at relatively high elevations, be- tween 1000 and 2500 m of altitude (Albouy and Causs- anel 1990). Most reports and descriptions indicate that C. aptera and C. pyrenaica are easily differentiated morphologically by the shape of male cerci (see Finot 1890; Azam 1901; Chopard 1922; Albouy and Causs- anel 1990). According to those authors, males of C. ap- tera have long, relatively thin and slightly curved cerci, while males of C. pyrenaica have short, broad and very curved cerci. In both species, cerci of females are short, thin and practically straight, with a slight curvature at the apex. However, Dohrn (1867), followed by Brunner von Wattenwy! (1882), Maccagno (1933) and Fontana et al (2021) considered that long, thin and slightly curved cerci together with short, broad and very curved cerci were part of the intraspecific variability of each taxon, and questioned the presence of C. aptera in the Pyrenees (or the presence of C. pyrenaica in the Alps). To compli- cate matters, as already noted by Maccagno (1933), the earliest descriptions of Pyrenean specimens of Chelidura as a differentiated taxon correspond to Forficula simplex Germar, 1825, which was described based on long cerci Pyrenean specimens (Germar 1825). Forficula simplex was subsequently included in the synonymy of C. aptera by Dohrn (1867), followed by Bolivar (1876), Brunner von Wattenwyll (1882), Finot (1890), Kirby (1904), Burr (1904), Sakai (1973), Harz and Kaltenbach (1976), and Herrera-Mesa (1999) among others, or treated as a vari- ety of C. aptera (Dubrony 1878). The known distribution range of Chelidura in the Pyr- enees is quite limited, with very few records in Andorra and the Spanish (Lapeira and Pascual 1980) and French slopes (Albouy and Caussanel 1990) (see “Species ac- counts” section). During field surveys aimed to document the persistence of the species in some of the classical lo- calities, we were surprised to find consistently specimens with long cerci (referred to as C. aptera in the literature; e.g. Lapeira and Pascual 1980; Albouy and Caussanel 1990), and short cerci (referred to as C. pyrenaica; op. cit.) coexisting at the same localities. These observations, together with the lack of consensus on the presence of C. aptera in the Pyrenees (see references above), prompt- dez.pensoft.net Pilar Jurado-Angulo et al.: Pyrenean species of Chelidura ed us to carry out a study to determine the correct iden- tification of long and short cerci specimens of Pyrenean Chelidura. For this purpose, we obtained cytochrome oxidase 1 (cox/) partial sequences of a few Pyrenean specimens, a representative of each cerci morphology, and we also raised under controlled conditions, a series of nymphs collected from the same clutch till they meta- morphose. The results of these analyses revealed that both long and short cerci males corresponded to a single taxonomic entity. With this main aim, the specific objectives of this work are: (1) to confirm the taxonomic identification of Chelidura specimens with long and short cerci present in the Pyrenees, (i1) discuss the taxonomic entity of the subspecies C. pyrenaica arverna from the French Massif Central, and (111) provide a species account including all known localities and synonymies of Pyrenean Chelidura. Material and methods Studied material, morphological study and distribution data Sampling was conducted in different localities of the Cata- lonian Pyrenees (Girona, Lleida), Andorra and Italy (Valle d’ Aosta). A total of 104 specimens, 95 specimens of C. pyrenaica and 9 specimens of C. aptera (see below) were collected. All specimens were collected by hand, photo- graphed in the field (when possible) and geo-referenced prior to being preserved in absolute ethanol, and then stored at —20 °C at the Museo Nacional de Ciencias Natu- rales (MNCN-CSIC) (Madrid, Spain). A set of 124 addi- tional specimens of C. aptera and C. pyrenaica from the MNCN-CSIC collection were used for the morphological study. A series of last instar nymphs from Tossa d’ Alp were maintained under controlled conditions until metamorpho- SIS, previous to preservation (11 males were obtained from nymphs). The 228 specimens studied are from: Chelidura pyrenaica (Gené, 1832): ANDORRA: Sant Julia de Loria: Bixessarri: Coll de la Gallina, 1933 m, 42°27'33.6"N, 1°27'03.7"E: 20-VI-2013, 2 females with eggs, M. Garcia-Paris, G. Garcia-Martin (MNCN_Ent 296001, 296015); La Rabassa, 1963 m, 42°26'21.7"N, 1°31'26.4"E: 20-VI-2013, 3 nymphs IV, 1 nymph V, 2 males, 5 females with eggs, M. Garcia-Paris, G. Garcia- Martin (MNCN_Ent 296016—296017, 269444, 296003— 296010). — FRANCE: Ariége: L’ Hospitalet-pres-l’ Andorre: 2 nymphs, | female, Dr. Martin (MNCN_Ent 283428— 283430). Hautes-Pyrénées: Bagneres-de-Bigorre: 20-[X- 1886, 1 male, 1 female, Collection A. Finot (MNCN_ Ent 7850, 283422). Pyrénées-Orientales: Canigou: 1 female, Col. Marquet (MNCN_ Ent 283425); 1 male, Masferrer (sub C. aptera) (MNCN_ Ent 7849); Coll d’Ares, 1508m, 42°21'58.7"N, 2°27'31.5"E: 17-VI-2013, 1 female, M. Garcia-Paris, G. Garcia-Martin (MNCN_Ent 295998), Mont-Louis: 1 male, 1 female, E. Simon leg., I. Bolivar det. (MNCN_ Ent 283426—283427). — Spain: Catalunya: Bar- Dtsch. Entomol. Z. 68 (2) 2021, 235-248 celona: Berga: Rasos de Peguera: 23-V-1991, 1 female, C. Martin (MNCN_Ent 122647); Montseny: 1 nymph, 3 males, 2 females, Masferrer(MNCN_Ent 122705, 122720- 122721, 122723—122725), Girona: Camprodon, 950 m: 15-[X-40, 5 males, J. Mat [J. Mateu] (MNCN_ Ent 122699, 122736—122739), 25-IX-40, 2 male, 2 females, J. Mat [J. Mateu] (MNCN_Ent 122697—122698, 122700—122701); La Molina: Tossa d’Alp, 2343 — 2484 m, 42°19'30.07"N, 1°54'10.89"E / 42°19'12.78"N, 1°53'45.57"E: 5-VII-2011, 22 nymphs, 8 females, 10 males, P. Pavon-Gozalo, M. Garcia-Paris, V. Salvador de Jesus (MNCN_ Ent 269465- 269466, 269468—269471, 269474, 269443, 269480- 269485, 296013—296014, 295972—295995): Puigcerda: 2 males, Zariquiey (MNCN_ Ent 122637, 122722); Puigmal, 2909 m: 1 male, Cazurro(MNCN_ Ent 122729); Setcases: Vallter, 1736 m, 42°24'11.50"N, 2°17'12.82"E: 4-VII-2011, 8 nymphs, P. Pavon-Gozalo, M. Garcia-Paris, V. Salvador de Jesus (MNCN_Ent 269460—269461, 269467, 269487, 295968—295971); 2174 m, 42°25'40.53"N, 2°15'58.56"E: 4-VII-2011, 23 nymphs, 2 males, 4 females, P. Pavon-Go- zalo, M. Garcia-Paris, V. Salvador de Jesus (MNCN_ Ent 269462-269464, 269472269473, 269442, 269475, 269477269479, 269486, 269489-269491, 295953- 295966, 13276); 2175 m, 42°25'36.7"N, 2°15'41.0"E: 17-VI-2013, 1 nymph IV, 1 nymph V, M. Garcia-Paris, G. Garcia Martin (MNCN_Ent 295999-296000); Toses: 26-[X-1932, 1 male, 1 female, A. Vilarrubia (MNCN_ Ent 122726—122727), Lleida: Bellver: 10-903 [X-1903], 1 Altitude [_J0 |_| 700 MS 1400 Mm 2100 ME 2800 23d male (MNCN_ Ent 122728); Caldes de Boi: VHI-1945, 3 males, Montada, (MNCN_Ent 122730—122732); Lles de la Cerdanya, 1935 m, 42°25'39.39"N, 1°39'51.73"E: 5-VI-2011, 4 nymphs, P. Pavon-Gozalo, M. Garcia- Paris, V. Salvador de Jesus (MNCN_ Ent 269476, 269488, 295996—295997):; Pto. Payas [Pallars]: Virgen de Arés [Alt Aneu]: 32 females, 29 males (MNCN_Ent 122638— 122644, 122646, 122648-122659, 122661—122677, 122679-122689, 122703, 122707—122718), 1923, 1 male, 2 females, M. Escalera (MNCN_Ent 122690, 122694— 122695), VIHI-1928, 4 females, 5 males, M. Escalera (MNCN_Ent 122660, 122678, 122702, 122704, 122706, 122691-122693, 122696); Salardu, 1.260 m: VIII-48, 1 nymph, E. Morales (MNCN_Ent 122645); Valle de Aran: Llenas: 1 nymph, 2 females (MNCN_Ent 122733-— 122735); Pirineos (without further indication): 1 male, Martorell (MNCN_Ent 122719); 1 female, 3 males, Col. Marquet (MNCN_ Ent 283423—283424, 283439283440) (specimens referenced from Pyrenees by Dubrony 1878 and Azam 1901) (Fig. 1). Chelidura aptera (Megerle, 1825): FRANCE: Savoie: Mont-Cenis: | female, H. Martin (MNCN_Ent 283431); Saint-Bernard [Col du Petit Saint Bernard]: 1 male, Brun- ner (MNCN_Ent 283438). — Itary: 3 females, 1 male, Durieu (MNCN_ Ent 283434—283437). Gressoney la T. [Trinité] (Piemonte, M. Rosa): VIII-935 [1935], 1 male, 1 female (C. Alzona) (MNCN_ Ent 283432—283433); Valle d’Aosta: Val Veny: Pré de Pascal, 1856 m, 45°48'20.2"N, Figure 1. Geographic distribution of Chelidura pyrenaica. Blue dots correspond to the species records, including both recent and old (see material and methods and species accounts for the localities). dez.pensoft.net 238 6°56'35.5"E: 28-VI-2012, 5 nymphs V, 2 females, 2 males, M. Garcia-Paris, G. Garcia-Martin (MNCN_ Ent 269452-269458, 296011—296012). Dry-mounted specimens were examined under a ste- reomicroscopy. 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 Fontana et al. (2002) with mi- nor modifications. The map depicting distribution range (Fig. 1) was made using the Natural Earth basemap, as implemented in QGIS vs. 3.8. DNA extraction and amplification Total DNA was obtained from six specimens (Table 1). DNA was extracted from one leg, using the DNeasy Blood and Tissue Isolation Kit (Qiagen, Hilden, Germa- ny), following the manufacturer’s instructions, and then stored at 4 °C until further processed. The polymerase chain reaction (PCR) consisted of, with occasional minor variations, 18.8 wL of distilled water, 2.5 uL of 10 x PCR buffer, 1 wL of dNTP mix (10 mM), 0.5 uL of MgCl2 (50 mM), 0.5 uL of each primer (10 uM), 0.2 wL of DNA polymerase (5u/uL) and 1 wL of DNA template, consist- ing of a final reaction volume of 25 wL. The universal pair of primers LCO1490 and HCO2198 (Folmer et al. 1994) were used to amplify a fragment of cox/, with the following PCR cycling profile: initial denaturation at 96 °C for 5 min, followed by 40 cycles at 94 °C for 30 s, 42 °C for 45 s and 72 °C for | min, and a final extension step at 72 °C for 5 min. After the amplification, 4 wL of the reaction was analyzed by electrophoresis on a 1% agarose gel. Samples with single bands were sent to the company Macrogen Inc. (Macrogen Europe, Madrid, Spain) for se- quencing in both directions. Pilar Jurado-Angulo et al.: Pyrenean species of Chelidura Phylogenetic analyses and species concept The cox] data set included four Pyrenean specimens (with diverse cerci morphology), two specimens from the Italian Alps, one specimen of C. p. arverna from Kirstova et al. (2020), and six additional specimens of C. aptera from Fontana et al. (2021) (Table 1). One additional spec- imen of Chelidurella vignai Galvagni, 1995 and another of Chelidurella thaleri Harz, 1980 (from Kirstova et al. 2020; Table 1) were used as closely related outgroups. We also included one specimen of Mesochelidura occidenta- lis Fernandes, 1973 and another of Anechura bipunctata (Fabricius, 1781) (from Kirstova et al. 2020; Table 1) as distant outgroups to root the phylogenetic analyses. Gen Bank accession numbers for the newly sequenced speci- mens are provided in Table 1. The obtained cox/ partial sequences were aligned with MAFFT v.7 (Katoh et al. 2019) using default pa- rameters. Uncorrected (p) pairwise genetic distances were estimated using PAUP* v.4.0a (Swofford 2002). The best substitution model obtained using Partition- Finder2 (Lanfear et al. 2016) was HKY +1 +y. An XML File was generated with BEAUti v.2.5.0 (Bouckaert et al. 2019) using a birth-death process model, and an un- correlated relaxed lognormal clock model under default parameters. Bayesian analyses were performed using in MrBayes v.3.2.6 (Ronquist et al. 2012) and BEAST v.2.6.3 (Bouckaert et al. 2019), through the CIPRES Science Gateway v.3 (Miller et al. 2010). The length of MCMC chain was 1,000,000 sampling every 1000. To check for convergence of the Markov chains Monte Car- lo (MCMC), posterior trace plots and effective sample sizes (ESS) were examined in TRACER v.1.7 (Rambaut et al. 2018). The first 25% of sampled trees were dis- carded, and using TreeAnnotator v.1.8.4 (Drummond et al. 2012), the results were summarized in a maximum clade credibility tree (MCC) and selecting a length of the nodes based on the median. Visualization and editing of the phylogenetic tree were carried out in FigTree v1.4.4 (Rambaut et al. 2018). Table 1. 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 Chelidura aptera MNCN Ent 296011 Italy: Valle d’Aosta: Val Veny: Pré de Pascal 45°48'20.2"N, 6°56'35.5"E MZ325323 Chelidura aptera MNCN Ent 296012 Italy: Valle d’Aosta: Val Veny: Pré de Pascal 45°48'20.2"N, 6°56'35.5"E MZ325324 Chelidura pyrenaica MNCN Ent 296013 Spain: Girona: La Molina: Tossa d’Alp 42°19'30.07"N, 1°54'10.89"E MZ325325 Chelidura pyrenaica MNCN Ent 296014 Spain: Girona: La Molina: Tossa d’Alp 42°19'30.07"N, 1°54'10.89"E MZ325326 Chelidura pyrenaica MNCN Ent 296015 Andorra: Sant Julia de Loria: Brxessarri: Coll de la Gallina 42°27'33.6"N, 1°27'03.7"E MZ325327 Chelidura pyrenaica MNCN Ent 296016 Andorra: Sant Julia de Loria: La Rabassa 42°26'21.7"N, 1°31'26.4"E MZ325328 Chelidura arverna France: Chalmazel (Kirstova et al. 2020) 45°40'33""N, 03°49'32"E MH853428 Chelidura aptera 5b-1 Switzerland, Valais, Col du Grand Saint-Bemard, Liddes, 2160 m 45°53'11.24"N, 7°11'24.35"E Fontana et al. (2021) Chelidura aptera 5b-5 Switzerland, Valais, Col du Grand Saint-Bemard, Liddes, 2160 m 45°53'11.24"N, 7°11'24.35"E Fontana et al. (2021) Chelidura aptera 2a-1 Italy, Piedmont (Biella), Pennine Alps, Lago del Mucrone, Oropa, 1910m = 45°37'43.54"N, 7°56'38.24"E Fontana et al. (2021) Chelidura aptera 2a-3 Italy, Piedmont (Biella), Pennine Alps, Lago del Mucrone, Oropa, 1910m = 45°37'43.54"N, 7°56'38.24"E Fontana et al. (2021) Chelidura aptera 17-1 Italy, Lombardy (Sondrio), Western Rhaetian Alps, Franscia, Lanzada, 1480m 46°17'21.4"N, 9°54'41.14"E Fontana et al. (2021) Chelidura aptera 18 Italy, Lombardy (Sondrio), Western Rhaetian Alps, Franscia, Lanzada, 1480m 46°17'21.4"N, 9°54'41.14"E Fontana et al. (2021) Chelidurella vignai Italy: Trento (Kirstova et al. 2020) 46°07'11"N, 11°15'40"E MH853430 Chelidurella thaleri Slovakia: Polana (Kirstova et al. 2020) 48°40'52''N, 19°30'29"E MH853433 Mesochelidura occidentalis Portugal: Monchique (Kirstova et al. 2020) 37°19'03"N, 08°35'18""W MH853427 Anechura bipunctata Mongolia: Ikh-tamir (Kirstova et al. 2020) 47°35'33"N, 101°12'60"E MH853426 dez.pensoft.net Dtsch. Entomol. Z. 68 (2) 2021, 235-248 Evolutionary (taxonomic) units within Chelidura were defined using the evolutionary species concept as dis- cussed in Sanchez-Vialas et al. (2020). The evolutionary Species concept considers species as “a single lineage of ancestral descendant populations of organisms that main- tain its identity from other such lineages and which has its own evolutionary tendencies and historical fate” (Wiley 1978, 1981; Wiley and Mayden 2000). Results Based on the phylogenetic analyses, studied specimens of Chelidura compose three well-supported clades (pos- terior probabilities = 1) (Fig. 2). A clade includes the Py- renean specimens (PP = 1), a second clade includes the Massif Central specimen, and the third clade includes the Alpine specimens (Valle d’ Aosta — Biella — Sondrio — Va- lais: Grand Saint Bernard) (PP = 1). The Alpine samples are geographically structured in two main subclades, one including samples from Valle d’ Aosta and Col du Grand Saint Bernard (PP = 0.99), the second from Biella and Sondrio (PP = 0.94). Uncorrected “p” distances between different groups based on cox/ partial sequences are sum- marized in Table 2. Pyrenean specimens (Andorra and Girona) form a monophyletic group of poorly differentiated sequences (uncorrected p ,..,,.. ranging from 0 to 0.03) (Fig. 2; Ta- ble 2). These samples include male specimens from Tossa d’Alp (Girona) and La Rabassa (Andorra) with typical short cerci (MNCN_ Ent 296014, 296016) and specimens from Tossa d’ Alp (Girona) with very long cerci (MNCN _ Ent 296013). The sister taxon relationship of the Alpine 239 clade with respect to the Pyrenean and Massif Central clades is poorly resolved (PP = 0.72) forming a possible polytomy with respect to Chelidurella. Genetic distance between the specimens of the Pyrenean and the Alpine clades is very large (uncorrected p ,...,. = 0.19-0.22) (Table 2). It is almost as high as those found among dif- ferent genera of Forficulidae (Table 2), suggesting that nucleotide changes in cox/ might be already saturated at that level. Male specimens included in the Pyrenean clade (Gi- rona and Andorra) present large variability in the shape of the cerci. Cerci range from long, almost straight con- vergent cerci (Figs 3A, 4A), to very curved, broad, short cerci (Fig. 3D). Short cerci present the maximum curva- ture at the middle, forming an angle of approximately 90°; cerci are wider at the base, strongly narrowed at the area of greatest curvature (Fig. 3D, E), and maintaining a more or less constant width up to the apex (Fig. 3D). Long cerci are sub-cylindrical, slightly curved at their maximum width near the base, progressively narrowed towards the apex, acuminate at the end (Fig. 3A). Cerci may present a ridge on the inner margin (Fig. 3B—F) or not (Fig. 3A). This ridge, when present, arises after the point of greatest curvature of the cerci and can continue to the apex of the cerci (Fig. 3D, E) or ending earlier, resembling a broad tooth (Fig. 3F). Intermediate speci- mens between these extreme shapes also occur (Figs 3B, C, F, 4C, D). In the same way, the diameter of the cerci is variable, including specimens with thick cerci compared to others with finer cerci. In all the individuals studied, cerci diameter expands over half or more of the width of the last segment. Males raised under controlled con- ditions from a single group of last instar nymphs from 1 Spain: Tossa d’Alp (short cerci) * Pa Spain: Tossa d’Alp (long cerci) * piraiees Chelidura pyrenaica Andorra: Coll de la Gallina (short cerci) * 1 : i) * 0.87 Andorra: La Rabassa (short cerci) Chelidura arverna Chelidura 0.72 Chelidura aptera 0.98 France: Chalmazel Massif Central 0.959 Italy: Valle d’Aosta * 0.99 Italy: Valle d'Aosta * Switzerland: Valais 4 0.616 Switzerland: Valais Alps 0.74 Italy: Biella 0.94 Italy: Biella 1 p Italy: Sondrio Italy: Sondrio Chelidurella vignai Chelidurella thaleri Mesochelidura occidentalis Anechura bipunctata Figure 2. Bayesian phylogenetic tree based on cox! partial sequences. The colours represent species and geographic areas: Pyrenees (blue), Massif Central (red) and Alps (green). Posterior probabilities are indicated for each clade. Sequences marked with an asterisk were obtained for this study, all other sequences were recovered from Kirstova et al. (2020) and Fontana et al. (2021). dez.pensoft.net 240 Pilar Jurado-Angulo et al.: Pyrenean species of Chelidura Table 2. Uncorrected (p) pairwise genetic distance matrix between specimens (short — long cerci) and taxa used in the phylogenetic analyses. Chelidura Chelidurella Chelidurella Anechura Mesochelidura $s aptera Cc. pyrenaica C. arverna vignai thaleri bipunctata occidentalis Valle Valais Biella Sondrio Long Short d’ Aosta cerci cerci Chelidura C. aptera 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 C. pyrenaica Long cerci 0.19-0.21 Short cerci 0.19-0.22 0.00-0.03 0.02-0.03 C. arverna 0.18-0.21 0.15 0.14-0.15 Chelidurella Ch. vignai 0.20-0.21 0.21 0.20-0.21 0.20 Ch. thaleri 0.20-0.21 0.19 0.19-0.20 0.21 0.15 Anechura A. bipunctata 0.22-0.23 0.22 0.22 0.22 0.20 0.20 Mesochelidura M. occidentalis 0.24-0.25 0.20 0.20 0.21 0.23 0.22 0.19 Figure 3. Cerci variation in Chelidura pyrenaica (Gené, 1832) males. Specimens from: (A) Puigcerda (Girona, Spain) (MNCN _ Ent_ 122637); (B) Pyrenees (Spain) (MNCN_Ent_ 122719); (C) Camprodon (Girona, Spain) (MNCN_Ent_ 122738); (D) Pyrenees (MNCN Ent 283424); (E) Caldes de Boi (Lleida, Spain) (MNCN_ Ent 122732); (F) Montseny (Barcelona, Spain) (MNCN _ Ent_ 122721). Scale bar = 1 mm. Tossa d’Alp present long, short and intermediate cerci (Fig. 4A, C, D). The specimens studied from the Alpine clade pres- ent long cerci with little curvature, cylindrical apically, progressively narrowed towards the apex and the inner margins without teeth or with one tooth. The diame- ter of the cerci of those specimens studied is general- ly smaller than that of the specimens of the Pyrenean clade. However, our sample is not representative of the dez.pensoft.net variability already reported for the Alpine clade (Burr 1912; Amiet 1961; David and Van Herrewege 1973; Sakai 1973; Caussanel and Albouy 1987; Albouy and Caussanel 1990; Herrera-Mesa 1999). Fontana et al. (2021) indicated that cerci variability in C. aptera is larger than previously considered, pending of a detailed geographic analysis. The Pyrenean clade is sister to the single sequence representing the Massif Central clade (PP = 0.87). The Dtsch. Entomol. Z. 68 (2) 2021, 235-248 C 241 D Figure 4. Live specimens of Chelidura pyrenaica (Gené, 1832) and cerci variation. A. Male from Tossa d’Alp (Girona, Spain) (MNCN Ent 296013). B. Female from Coll de La Rabassa (Andorra). C. Male from Tossa d’Alp (Girona, Spain) (MNCN_ Ent 269481). D. Male from Tossa d’ Alp (Girona, Spain). Photographs ex situ by M. G.-P. genetic distance between Pyrenean and Massif Central populations is quite large (uncorrected p ,..., = 0.14- 0.15). Among the large series of Pyrenean specimens studied we did not find the cerci morphology described for C. p. arverna by David & Van Herrewege (1973), Kirstova et al. (2020) and Fontana et al (2021). Cerci of the specimen included in the Massif Central clade, from Chalmazel (France), are more robust, relatively wider and less curved than cerci of the specimens of the Pyrenean clade. David and Van Herrewege (1973) confirmed that the morphology of the Pyrenean populations and those of the Massif Central (morphometric traits and male cerc1), differ statistically. Male genitalia from specimens of the Pyrenees, Al- pine and Massif Central clades, including the lectotype of C. pyrenaica, the neotype of C. aptera and the holo- type of C. p. arverna, were studied in detail and photo- graphed by Fontana (1999) and Fontana et al. (2021). Maccagno (1933) also provided an illustration of the male genitalia of Pyrenean specimens corresponding to C. pyrenaica. The genitalia of the male specimens of C. pyrenaica we examined (Virgen de Ares, Lleida) match the description presented by Maccagno (1933) and Fontana (1999); variability is however large, including size of parameres. They differ from those of the Alpine specimens, by showing thinner parameres with almost parallel margins (shorter and curved on the external mar- gin in Alpine specimens). Male genitalia of typical C. pyrenaica and the holotype of C. p. arverna do not differ significantly, although C. p. arverna seems to present a more arcuate vesicle. Species accounts Chelidura arverna David & Van Herrewege, 1973 Stat. nov. Chelidura pyrenaica arverna David & Van Herrewege, 1973: 40. Terra typica: «Massif Central: Mont Mézenc». Holotype at the Muséum d’ Histoire naturelle de Paris (David & Van Herrewege 1973). Albouy & Caussanel (1990: 180) wrote the species name as “C. p. averna’. Published records. FRANCE: Cantal (Chopard 1922 sub C. aptera, Chopard 1951 sub C. aptera; Sakai 1973 sub C. aptera;, Harz and Kaltenbach 1976 sub C. aptera); Le Lioran (Burr 1904 sub C. aptera),; Le Lioran, prairies voisines de la station (Finot 1890 sub C. aptera). Haute- dez.pensoft.net 242 Loire: Massif Central (Amiet 1961 sub C. aptera; Causs- anel and Albouy 1987 sub C. aptera and C. pyrenaica;, Albouy and Caussanel 1990 sub C. aptera and C. pyrena- ica, Herrera-Mesa 1999 sub C. pyrenaica), Mont Mézenc (David and Van Herrewege 1973; Albouy and Caussanel 1990 sub C. pyrenaica spp. averna [arverna]). Loire: Chalmazel: Station de Chalmazel, 1137 m, 45°40'33"N, 3°49'32"E (Fontana et al. 2021). Lozére (Chopard 1922 sub C. aptera; Chopard 1951 sub C. aptera; Sakai 1973 sub C. aptera; Harz and Kaltenbach 1976 sub C. aptera). Puy-de-Doéme (Chopard 1922 sub C. aptera; Chopard 1951 sub C. aptera; Sakai 1973; Harz and Kaltenbach 1976 sub C. aptera): Mont Doré (Fontana et al. 2021). Dubious assignment: FRANCE: Ardéche (Harz and Kaltenbach 1976 sub C. aptera): Astet (Chopard 1951 sub C. aptera). Chelidura pyrenaica (Gené, 1832) Forficula simplex Germar, 1825: pl. 17 (nomen oblitum) syn. nov. Terra typica: “... in Pyrenaeis...”. Forficula pyrenaica Gené, 1832: 227 (nomen protectum). Terra typica: “__Pirenei...”. Lectotype designated by Fontana (1999) (male, speci- men number 2363 at MRSNT). The neotype designation mentioned by Harz and Kaltenbach (1976) with specimens from “Riba Freser” (sic), is not valid (Fontana 1999). Forficula dilatata Burmeister, 1838: 755. Terra typica “In den Pyrenaen” ? Forficula pyrenaea Herrich-Schaffer, 1840: 31. Terra typica not indi- cated. A synonym of either C. pyrenaica or Pseudochelidura sinuata (Germar, 1825) (Herrich-Schaffer 1840). Chelidura dilatata (Burmeister, 1838): Brunner von Wattenwyll 1882: 25 Chelidura pyrenaica (Gené, 1832): de Bormans and Krauss 1900: 108. Sakai (1973: 175) wrote by mistake «Chelidura pyrenaticay. Published records. ANDORRA (David and Van Herre- wege 1973; Sternmann 1981 sub C. aptera). — FRANCE: Ariége (Dubrony 1878 sub C. aptera; Finot 1890 sub C. dilatata, Azam 1901 sub C. dilatata,; Chopard 1951; Sakai 1973): 1500 m (Chopard 1922); Anglade [Cirque d’Anglade] (David and Van Herrewege 1973); L’Hos- pitalet-pres-Il’ Andorre (Dusoulier 2004); Montagnes de VAri¢ge (Marquet 1877 sub C. dilatata); Sollau [not found] (David and Van Herrewege 1973). Haute-Ga- ronne (Marquet 1877 sub C. dilatata): Vallé d’Esquieres Luchon bei 1400 m [Bagneres-de-Luchon] (Fieber 1853 sub F) dilatata). Hautes-Pyrénées (Dubrony 1878 sub C. aptera,; Brunner von Wattenwy! 1882 sub C. dilatata, Chopard 1951 sub C. aptera; Sakai 1973): au-dessus de 1500 m (Chopard 1922); 4 une hauteur de 2000 a 2500 m (Finot 1890 sub C. dilatata); a partir de 1000 m (Chopard 1922 sub C. aptera), Bagneres-de-Bigorre (Azam 1901 sub C. dilatata, Chopard 1951); Bagneres-de-Bigorre, au lac Bleu (Azam 1901 sub C. dilatata); environs de Ba- eneres-de-Bigorre (Finot 1890 sub C. dilatata); Glacier de Neouvielle (Burr 1912 sub C. dilatata, Sakai 1973); Pic de Nere (David and Van Herrewege 1973); Pic-du- Midi (Finot 1890 sub C. dilatata; Chopard 1951); Pic du dez.pensoft.net Pilar Jurado-Angulo et al.: Pyrenean species of Chelidura Midi de Big 2000-2800 m (Harz and Kaltenbach 1976 sub C. aptera),; Saint-Lary [Saint-Lary-Soulan] (Dau- phin 1987); Seincourt [not found] (David and Van Her- rewege 1973). Pyrénées-Atlantiques (Albouy and Causs- anel 1990). Pyrénées-Orientales (Dubrony 1878 sub C. aptera, Finot 1890 sub C. dilatata; Azam 1901 sub C. dilatata, Chopard 1922): au-dessus de 1500 m (Cho- pard 1922); Canig6 (Finot 1890 sub C. dilatata, Azam 1901 sub C. dilatata; Borelli 1905 sub C. dilatata, Burr 1912 sub C. dilatata, Amiet 1961; Sakai 1973; David and Van Herrewege 1973); Canigou, a partir de 1000 metres (Xambeu 1907 sub C. dilatata, Chopard 1951); Canig6, a partir de 1200 m d’altitude, jusqu’a 2400 (Xambeu 1903 sub C. dilatata),; Coubezet (Xambeu 1907 sub C. dila- tata); Font-Romeu [Font-Romeu-Odeillo-Via] (Borelli 1905 sub C. dilatata, Sakai 1973; Vancassel and Foraste 1980; Vancassel 1984); Le Vernet [Vernet-les-Bains] (Dubrony 1878 sub C. aptera; Azam 1901 sub C. dila- tata), Le Vernet [Vernet-les-Bains], pres Prades (Finot 1890 sub C. dilatata,; Chopard 1951); Mont-Louis (Amiet 1961; Steinmann 1981 sub C. aptera); Rouquette [Pic de la Rouquette] a partir de 1200 m d’altitude, jusqu’a 2400 (Xambeu 1903 sub C. dilatata); Thues, Haute vallée de la Carang¢a (Hamon 1956); Val d’Eyne (Chopard 1951; Hamon 1956); Vallée supérieure du Tech (Borelli 1905 sub C. dilatata,; Sakai 1973). Pyrénées (département not indicated) (Serville 1839 sub F’ aptera and F. simplex; Fieber 1853 sub F. simplex and F: dilatata; Dubrony 1878 sub C. aptera and C. a. var. simplex; Brunner von Watten- wyl 1882 sub C. dilatata; de Bormans and Krauss 1900; Azam 1901 sub C. aptera; Kirby 1904; Borelli 1905 sub C. dilatata, Amiet 1961; David and Van Herrewege 1973; Caussanel and Albouy 1987; Albouy and Caussanel 1990): les parties élevées (Azam 1901 sub C. dilatata), localités élevées (Houlbert 1900 sub C. dilatata); south- ern Europe from Pyrenees and Southern France (Sakai 1973 sub C. aptera). — SPAIN (province not indicated): Espagne en montagne, entre 1000 et 2500 m d’altitude (Albouy and Caussanel 1990 sub C. aptera); Norte de Espafia (Cazurro Ruiz 1888 sub C. dilatata). Aragon: Huesca: Coll de Basibé, 2000-2200 m (Borelli 1926; Sakai 1973); Hospital de Benasque, Maladeta, 1775 m (Borelli 1926; Sakai 1973); Valibierne-Tal bei Benasque, 2000-2400 m (Borelli 1926; Sakai 1973). Catalunya: Barcelona (Herrera-Mesa 1999): Espinalbet (Lapei- ra and Pascual 1980); Montseny (Lapeira and Pascual 1980). Girona (Herrera-Mesa 1999): Camprodon (Ca- zurro Ruiz 1888 sub C. dilatata, Novellas 1901); Cam- prodén 950 m (Lapeira and Pascual 1980); Camprodon (“a native of the upper regions of the Pyrenees, where it occurs at an elevation of 6000ft.—8000ft.”) (Burr 1904); Col de Tosas [Collada de Toses] (David and Van Herre- wege 1973); Nuria [Vall de Nuria] (Lapeira and Pascual 1980); Nuria, pinar de la Virgen, a mas de 2000 metros (Navas 1921); Puigcerda (Lapeira and Pascual 1980); Puigmal [Puigmal d’Er], 2909 m (Lapeira and Pascual 1980); Riba Freser [Ribes de Freser] (Harz and Kalten- bach 1976; Albouy and Caussanel 1990; Fontana 1999); Dtsch. Entomol. Z. 68 (2) 2021, 235-248 Figure 5. Live specimens of Chelidura pyrenaica (Gené, 1832) from Andorra and typical habitat. A. Female with eggs from Coll de la Gallina (Andorra). B. Early instar nymph from Coll de La Rabassa (Andorra). C. Late instar nymph from Coll de la Rabassa (Andorra). D. Typical habitat where C. pyrenaica complete its development (Coll de la Gallina, Andorra; June). E. Slopes of Tossa d’ Alp (Girona; July) where specimens of C. pyrenaica showing a wide variability of cerci shape coexist. Photographs by M. G.-P. Ripolles: Toses [Collada de Toses] (Lapeira and Pascual 1980); Riu (BVdb 2021); Ull de Ter [Ulldeter] (Lapeira and Pascual 1980). Lleida (Herrera-Mesa 1999): Aransa [Aranser] (BVdb 2021); Bellver [Bellver de Cerdanya]| (Lapeira and Pascual 1980); Bellver de Cerdanya (BVdb 2021); Bor (BVdb 2021); Caldas Bohi [Caldes de Boi] (Lapeira and Pascual 1980); Martinet (Boeseman 1954; Sakai 1973; Sakai 1973 sub C. aptera), Parque Nacio- dez.pensoft.net 244 nal de Aigues Tortes (Balcells et al. 1962); Pto. Payas [Pallars]: Virgen de Ares [Alt Aneu] (Lapeira and Pas- cual 1980); Tirvia (BVdb 2021); Tornafort (BVdb 2021); Valle de Aran [Val d’Aran] (Lapeira and Pascual 1980); Val d’Aran: Port de Viella (Borelli 1926; Sakai 1973; Lapeira and Pascual 1980); Val d’Aran: Salardt, 1260 m (Lapeira and Pascual 1980). Pirineos (provincia not indicated) (Fischer 1853 sub C. dilatata, Bolivar 1878 sub C. aptera; Martorell Pefia 1879; Cazurro Ruiz 1888 sub C. aptera and C. dilatata,; Burr 1910; Popham 1968; Sakai 1973; Harz and Kaltenbach 1976; Lapeira and Pas- cual 1980; Caussanel and Albouy 1987; Steinmann 1989; Fontana 1999; Herrera-Mesa 1999; Guillet and Vancassel 2001; Kirstova et al. 2020 sub Chelidura; Fontana et al. 2021) (Fig. 1). Fontana et al. (2021: fig. 13) commented on a spec- imen morphologically assignable to C. pyrenaica from the Sierra Nevada Mountains in Southern Spain (Picacho de Veleta; Museum National d’ Histoire Naturelle, Paris). As Fontana et al. (2021) discussed, it is quite possible that the specimen could be mislabelled, as it has already happened with other specimens of Dermaptera labelled erroneously from the Sierra Nevada Mountains, other- wise a quite well explored mountain chain (Garcia-Paris 2017). The presence of Chelidura in the Sierra Nevada Mountains should be treated as doubtful until additional specimens come to light. Notes on Natural History Chelidura pyrenaica is found in mountain slopes, be- tween 1000 and 2500 m, usually in pastures in areas cov- ered by flat stones, near the forest edge or in open areas (Fig. 5D, E) (Borelli 1905; Chopard 1922; Chopard 1951; David and Van Herrewege 1973; Albouy and Caussanel 1990). The geologic substrates of the area are diverse and complex, dominated by schists and limestone (see for a general overview Dendaletche 1982). Adult specimens are usually found under stones, bark of fallen trees and clods of earth in summer and fall (Azam 1901; Xambeu 1903; Chopard 1922; Albouy and Caussanel 1990). Xam- beu (1903, 1907) and Chopard (1951) mentioned that C. pyrenaica can be also found in spring. Mating takes place in April or May, in galleries that earwigs dig under their shelters. Females lay the eggs grouped in a shallow excavation at the end of one of these galleries and take care of them until the larvae hatch (Xambeu 1903; Causs- anel and Albouy 1987; Albouy and Caussanel 1990). We observed female attending eggs in the second half of June in Andorra (Fig. 5A). Upon disturbance females hide in small burrows, but usually return soon to the egg mass and start moving the eggs to the burrow, holding them one by one with their mandibles. Herter (1943) indicates that females may lay 40-45 eggs per clutch. Diverse nymphal stages were observed in the second half of June in Andorra and in the first half of July in Coll d’ Ares (Gi- rona) also attended by females (Fig. 5B, C). dez.pensoft.net Pilar Jurado-Angulo et al.: Pyrenean species of Chelidura Discussion 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 ex- ample MNCN_ Ent 296013; Fig 4A), short, and interme- diate cerc1 (MNCN_ Ent 269481; Fig. 4C) also support that cerci variability corresponds to a single taxon. Populations of Chelidura from the French Massif Central have been treated as a differentiated subspecies, C. pyrenaica arverna (David and Van Herrewege 1973; Kirstova et al. 2020). The large genetic distance observed between Pyrenean and Massif Central populations of Che- lidura suggests that that they have been isolated for long time. Lasting isolation between the Pyrenean and Central Massif populations is also supported by the morpholog- ical differentiation observed in male cerci. Reciprocal monophyly, long isolation reflected by large sequence di- vergence, and cerci morphological differentiation at mor- phometric level, suggest that C. arverna likely represents a separate taxonomic unit with respect to C. pyrenaica as previously suggested by Fontana et al. (2021). Using the evolutionary species concept (Wiley 1978, 1981; Wiley and Mayden 2000), there is little doubt that the Massif Central populations of Chelidura can be considered to represent an independent species: Chelidura arverna Da- vid and Van Herrewege, 1973 stat. nov. Intraspecific variability of morphological structures, as pygidium or cerci, is well known in earwigs (Srivastava 1970; Simpson and Mayer 1990; Tomkins and Simmons 1996; Garcia-Paris 2017; Kirstova et al. 2020). Many spe- cies of Dermaptera show large variability in the size and shape of male cerci (Dohrn 1867; Diakonov 1925; Ollason 1970; Srivastava 1970; Mourier 1986; Simpson and May- er 1990; Gonzalez-Miguens et al. 2020; Garcia-Paris et al. in press). The level of variability found in Pyrenean Che- lidura is apparently higher than the levels of variability found by these previous authors in other taxa (Fontana et al. 2021; Fig. 3). This large variability in male sexual char- acters might be a consequence of strong sexual selection (Kawano 2006; Brown 2007). Alternatively, the large vari- ability observed could be a consequence of the absence of directional selective pressures as Kirstova et al. (2020) mentioned as a possible explanation for the variability of the shape of the pygidium in some species of Chelidurella. The taxonomic implications of the large shape vari- ability in male cerci need to be addressed in the case of Pyrenean Chelidura. The presence of specimens of C. pyrenaica in the Pyrenees with long cerci was already mentioned by Borelli (1905), who said: “Parmi les in- Dtsch. Entomol. Z. 68 (2) 2021, 235-248 dividus trouvés sur les flancs du Canigou, trois ont les branches de la pince tres allongées, légerement arquées, ne se touchant pas a l’extrémité et pourvues en dedans, vers le milieu, d’une petite dent a peine visible...”. Dohrn (1867) also mentioned that he had seen specimens of C. pyrenaica (sub C. dilatata) with long narrow cerci not typical for this species. More recently, Maccagno (1933) and Fontana et al. (2021), based on examination of male genitalia made a clear statement indicating that Pyrenean specimens with long cerci corresponded to C. pyrenaica. But many other authors disregarded these considerations treating long cerci Pyrenean specimens as C. aptera, and consequently reporting the presence of this species in the Pyrenees (Serville 1831, 1839; Fieber 1953; Bolivar 1878; Dubrony 1878; Cazurro Ruiz 1888; Azam 1901; Chopard 1922; Chopard 1951; Boeseman 1954; Amiet 1961; Sakai 1973; Harz and Kaltenbach 1976; Lapeira and Pascual 1980; Steinmann 1981; Caussanel and Albouy 1987; Al- bouy and Caussanel 1990; Herrera-Mesa 1999). Most of the confusion derived from the early syn- onymization of Forficula simplex Germar, 1825, de- scribed based on Pyrenean specimens displaying long cerci (Germar, 1825), with C. aptera (Dohrn 1867; Boli- var 1876; Brunner von Wattenwyll 1882; Finot 1890; Kirby 1904; Burr 1904; Sakai 1973; Harz and Kaltenbach 1976; Herrera-Mesa 1999). The treatment of F’ simplex as a synonym of C. aptera carried two consequences, first the early inclusion of the Pyrenees within the geographic range of C. aptera, leading to the current confusion, and second, the unnecessary descriptions of Forficula pyrena- ica Gené, 1832 and Forficula dilatata Burmeister, 1838, based on specimens also from the Pyrenees. According to the principle of priority (article 23 of International Code of Zoological Nomenclature, ICZN 1999), the name Forficula simplex Germar, 1825 has nomenclatural prior- ity over Forficula pyrenaica Gené, 1832. However, both names meet the provisions of the article 23.9.1 (ICZN, 1999) to retain prevailing usage in order to assure nomen- clatural stability. In consequence, we propose the rever- sion of precedence of F: simplex with respect to F’ pyrena- ica. Forficula simplex Germar, 1825 has not been used as a valid species name after 1899, and Forficula pyrenaica Gené, 1832 has been used in in at least 25 works, pub- lished by at least 10 authors in the immediately preceding 50 years and encompassing a span of not less than 10 years (see for example David and Van Herrewege 1973; Sakai 1973; Harz and Kaltenbach 1976; Lapeira and Pascual 1980; Vancassel and Foraste 1980; Shah 1984: Vancassel 1984; Caussanel and Albouy 1987; Dauphin 1987; Pascual 1988; Steinmann 1989, 1993; Albouy and Caussanel 1990; Dendaletche 1982; Coutin 1983; Plate 1987; Haas 1995; Herrera-Mesa 1996, 1999; Guillet and Vancassel 2001; Klass 2001; Barrientos 2004; Dusoulier 2004; Matzke and Klass 2005; Costa 2006; Leraut 2007; Kirstova et al. 2020; Fontana et al. 2021). Therefore, and according to the Article 23.9.2. of the International Code of Zoological Nomenclature (ICZN 1999), the name Forficula pyrenaica Gené, 1832 is considered a nomen 245 protectum with nomenclatural precedence over the name Forficula simplex Germar, 1825, a nomen oblitum. Chelidura pyrenaica has been recorded in the Alps (Burr 1912; Amiet 1961; David and Van Herrewege 1973; Sakai 1973; Caussanel and Albouy 1987; Albouy and Caussanel 1990; Herrera-Mesa 1999) and C. aptera in the Massif Central (Finot 1890; Burr 1904; Chopard 1922; Chopard 1951; Amiet 1961; Sakai 1973; Harz and Kaltenbach 1976; Caussanel and Albouy 1987; Al- bouy and Caussanel 1990), but we conclude, totally in agreement with Fontana et al. (2021), that the reports of C. pyrenaica from the Alps, and those of C. aptera in the Massif Central, should be disregarded, and assigned to C. aptera and C. arverna respectively. Acknowledgements We thank Pilar Pavon Gozalo, Vladimir Salvador de Jesus and Gonzalo Garcia for their help during field surveys. This work was possible thanks to Mercedes Paris, curator of Entomology of the Museo Nacional de Ciencias Na- turales (Madrid), who helped us during the revision of the dry-preserved materials from the Entomological Col- lection and for assistance while taking images. We thank Paolo Fontana, Jose Luis Ruiz, and an additional anon- ymous reviewer for relevant suggestions that improved this manuscript. We also thank Markéta Kirstova and Petr Koéarek for comments and suggestions on this project. Thanks to Alberto Sanchez Vialas for help with the distri- bution map and to Fernando Garcia Guerrero for help with the photographs of the male genitalia. This work was par- tially funded by the project grant PID2019-110243GB-100 / AEI/10.13039/501100011033 (Ministerio de Ciencia, In- novacion y Universidades, Spain) to MG-P. References Albouy V, Caussanel C (1990) Dermapteres et perce-oreilles. Faune de France n° 75. Fédération Francaise des Sociétés de Sciences Na- turelles, Paris, 245 pp. Amiet JL (1961) Chelidura pyrenaica Bon. Dans les Alpes savoyardes (Dermaptere). L’Entomologiste 17(3): 33-35. Azam J (1901) Catalogue synonymique et systématique des Orthoptéres de France. Miscellanea Entomologica 9: 17-32. Balcells Rocamora E, Espafiol F, Montserrat P, Selga D (1962) Avance del estudio del Parque Nacional de Aigties Tortes. Actas del Tercer Congreso Internacional de Estudios Pirenaicos: Gerona, 1958, vol. 2, seccion 2: Climatologia, edafologia, botanica y zoologia. Instituto de Estudios Pirenaicos, Zaragoza. Barrientos JA (2004) Curso practico de Entomologia. Manuals de la Universitat Autonoma de Barcelona. 41. Entomologia. Asociacion Espafiola de Entomologia, CIBIO-Centro Iberoamericano de Bio- diversidad & Universitat Autonoma de Barcelona, 947 pp. ISBN: 84-490-2383-1. Boeseman M (1954) The Dermaptera in the Museums at Leiden and Amsterdam. Zoologische Verhandelingen 21: 1-122. dez.pensoft.net 246 Bolivar I (1878) Sindpsis de los ortopteros de Espafia y Portugal. Im- prenta de T. Fortanet, Madrid, 333 pp. [+ 7 pls.] Borelli A (1905) Sur quelques Forficules des Pyrénées. La Feuille des Jeunes Naturalistes, série IV 35: 49-51. Borelli A (1926) Forficuliden aus dem nordlichen und ostlichen Span- ien, gesammelt von Dr. F. Haas in den Jahren 1914 — 1919 und von Prof. Dr. A. Seitz im Jahre 1923. Senckenbergiana 8: 166-167. Bouckaert R, Vaughan TG, Barido-Sottani J, Duchéne S, Fourment M, Gavryushkina A, Heled J, Jones G, Kthnert D, De Maio N, Matschiner M, Mendes F, Miller N, Ogilvie H, du Plessis L, Pop- inga A, Rambaut A, Rasmussen D, Siveroni I, Suchard M, Wu C, Xie D, Zhang C, Stadler T, Drummond A (2019) BEAST 2.5: An advanced software platform for Bayesian evolutionary analysis. PLoS Computational Biology 15(4). https://doi.org/10.1371/jour- nal.pcbi. 1006650 Brown GS (2007) Sperm competition and male forceps dimorphism in the European earwig Forficula auricularia (Dermaptera: Forficu- lina). PhD Thesis, University of St Andrews, Scotland. http://hdl. handle.net/10023/409 Brunner von Wattenwyl C (1882) Prodromus der européischen Ortho- pteren. von Wilhelm Engelmann, Leipzig, 466 pp. Burmeister HCC (1838) Handbuch der Entomologie. Enslin, Berlin, 2, 756 pp. Burr M (1904) Synopsis of the Orthoptera of Western Europe. The En- tomologist’s Record and Journal of Variation 16: 42-44. Burr M (1910) 17. Orthoptera. Dermatoptera. [Pp. 1-12 [93—104]]. In: Sjostedt Y (Ed.) Wissenschaftliche Ergebnisse der Schwedischen Zoologischen Expedition nach dem Kilimandjaro, dem Meru und den umgebenden Massaisteppen Deutsch-Ostafrikas 1905-1906 (Vol. 3). Palmquists Aktiebolag, Stockholm, 636 pp. [+ 37 pls.] Burr M (1912) Die Dermapteren des k.k. naturhistorischen Hofmuse- ums in Wien. Annalen des Naturhistorischen Museums in Wien 26: 63-108. https://www.biodiversitylibrary.org/page/4460789 BVdb (Biodiversidad Virtual Database) (2021) Insectarium virtual. Ge- nus Chelidura. https://www.biodiversidadvirtual.org/insectarium/ Chelidura-Latreille-1825-cat3306.html [last accessed 20 Jan. 2021] Caussanel C, Albouy V (1987) Biosystématique des Dermapteéres de France. Annales de la Société Entomologique de France 23: 77-98. Cazurro Ruiz M (1888) Enumeracion de los ortopteros de Espafia y Portugal. Anales de la Sociedad Espafiola de Historia Natural 17: 435-513. Chopard L (1922) Orthopteres et Dermaptéres. Fauna de France, 3. Paul Lechevalier, Paris, 212 pp. Chopard L (1951) Orthoptéroides. Faune de France, 56. Paul Lecheva- lier, Paris, 359 pp. Costa JT (2006) The other insect societies. Harvard University Press, Cambridge, Mass., London, 812 pp. Coutin R (1983) Caractéres originaux de la faune entomologique du Queyras calcaire (Hautes-Alpes). Comptes rendus du 108e Congrés national des sociétés savantes, Grenoble, 1983, Section des Sciences 1(2): 247-255. [C.THS., Paris] Dauphin P (1987) A propos de quelques captures de Dermapteéres. Bul- letin de la Société Linnéenne de Bordeaux 15(1): 27-29. David J, Van Herrewege C (1973) Une sous-espéce frangaise de Che- lidura pyrenaica (Bonelli) (Dermapteres): C. pyrenaica arverna, ssp. n. Bulletin de la Société Linnéenne de Lyon 42(10): 31-41. https://doi.org/10.3406/linly. 1973.10029 dez.pensoft.net Pilar Jurado-Angulo et al.: Pyrenean species of Chelidura de Bormans A, Krauss H (1900) Forficulidae und Hemimeridae. Das Tierreich 11, von R. Friedlander und Sohn, Berlin, 142 pp. https:// www. biodiversitylibrary.org/page/43 101643 Dendaletche C (1982) Guia de Los Pirineos. Omega, Barcelona, 790 pp. [Originally published as: Guide du naturaliste dans les Pyrenées Oc- cidentales. Delachaux & Niestlé, Neuchatel] Diakonov DM (1925) Experimental and biometrical investigations on dimorphic variability of Forficula. Journal of Genetics 15: 201-232. https://doi.org/10.1007/BF02983 107 Dohrn H (1867) Versuch einer Monographie der Dermapteren. Ento- mologische Zeitung Herausgegeben von dem entomologischen Vereine zu Stettin 28: 341-343. Drummond AJ, Suchard MA, Xie D, Rambaut A (2012) Bayesian phy- logenetics with BEAUti and the BEAST 1.7. Molecular Biology and Evolution 29: 1969-1973. https://doi.org/10.1093/molbev/mss075 Dubrony A (1878) Liste des Orthopteres recueillis jusqu’ici en Ligurie. Annali del Museo Civico di Storia Naturale di Genova 12: 5—25. Dusoulier F (2004) Les Odonates et les Orthoptéres de la collection de Jean Des Abbayes (1900-1975): éléments du biopatrimoine vendeéen. Le naturaliste Vendéen 4: 57-64. Fieber FX (1853) Synopsis der europadischen Orthopteren. Lotos, Zeitschrift fiir Naturwissenschaften 3: 252-258. Finot A (1890) Faune de la France. Insectes. Orthopteres: Thysanoures et Orthoptéres proprement dits. Deyrolles, Paris, 322 pp. https:// www. biodiversitylibrary.org/page/8 185851 Fischer LH (1853) Conspectus Systematicus Orthopterorum Europae. Engelmann, Lipsiae, 16 pp. Folmer O, Black M, Hoeh W, Lutz R, Vrijenhoek R (1994) DNA prim- ers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology 3: 294-299. Fontana P (1999) The type specimens of Forficula described by Gené in 1832, at present belonging to the genus Chelidura Latreille, 1825 (Insecta Dermaptera). Bollettino del Museo Regionale di Scienze Naturali Torino 16: 127—150. Fontana P, Buzzetti FM, Cogo A, Odé B (2002) Guida al riconosci- mento e allo studio di Cavallette, Grilli, Mantidi e Insetti affini del Veneto. Ed. Museo Naturalistico Archeologico di Vicenza, 592 pp. Fontana P, Pedrazzoli P, Malagnini V, Ruzzier E, Marangoni F, Kocarek P (2021) Toward a revision of the genus Chelidura Latreille, 1825: designation of the Neotype for Chelidura aptera (Megerle in Char- pentier, 1825) (Dermaptera: Forficulidae). Memorie della Societa Entomologica Italiana 97: 279-302. [pISSN 0037-8747, eISSN 2282-1228] https://doi.org/10.408 1/memoriesei.2020.279 Garcia-Paris M (2017) Taxonomy of Iberian Anisolabididae (Dermap- tera). Acta Zoologica Academiae Scientiarum Hungaricae 63(1): 29-43. https://doi.org/10.17109/AZH.63.1.29.2017 Garcia-Paris M, Jurado-Angulo P, Martinez-Pérez S, Mico E (submit- ted) Rediscovery of Forficula iberica Steinmann, 1981 (Dermap- tera: Forficulidae). Zootaxa. Gené G (1832) Saggio di una Monografia delle Forficule indigene. An- nali delle Scienze del Regno Lombardo-Veneto 2: 215-228. Germar EF (1825) Fauna insectorum Europae. Fasciculus 11. Kimmel, Halae, 25 pls. Gonzalez-Miguéns R, Mujfioz-Nozal E, Jiménez-Ruiz Y, Mas-Peinado P, Ghanavi HR, Garcia-Paris M (2020) Speciation patterns in the Forficula auricularia species complex: cryptic and not so cryptic Dtsch. Entomol. Z. 68 (2) 2021, 235-248 taxa across the western Palaearctic region. Zoological Journal of the Linnean Society 190(3): 788-823. https://doi.org/10.1093/zoolinne- an/zlaa070 Guillet S, Vancassel M (2001) Dermapteran life-history evolution and phylogeny with special reference to the Forficulidae family. Evolu- tionary Ecology Research 3(4): 477-486. Haas F (1995) The phylogeny of the Forficulina, a suborder of the Dermaptera. Systematic Entomology 20(2): 85-98. https://doi. org/10.1111/).1365-3113.1995 tb00085.x Hamon J (1956) Notes de chasse sur les Orthoptéroides récoltés dans les Pyrénées-Orientales et dans les régions limitrophes déterminés par M. Broquet. Documents faunistiques et écologiques. Vie et Milieu, Observatoire Océanologique — Laboratoire Arago 7(3): 418-422. Harz K, Kaltenbach A (1976) Die Orthopteren Europas III — The Or- thoptera of Europe HI. W. Junk, The Hague 12, 434 pp. https://doi. org/10.1007/978-94-017-25 13-2 Herrera-Mesa L (1996) A preview of the catalogue of the Dermaptera of Spain. Taxonomy of the Dermaptera. Proceedings, XX International Congress of Entomology, Firenze, Italy, August 27, 1996. Tokyo, Sakai, 426 pp. Herrera-Mesa L (1999) Catalogue of the Dermaptera of Spain. New- book Ed., Navarra, 203 pp. Herrich-Schaffer GA (1840) Nomenclator entomologicus: Verzeichniss der europaischen Insecten; zur Erleichterung des Tauschverkehrs mit Preisen versehen. Zweites Heft. Coleoptera, Orthoptera, Dermatop- tera und Hymenoptera. Pustet, Regensburg, 2, 244 pp. [+ 8 pls.] Herter K (1943) Zur fortpflanzungsbiologie eines lebendgebarenden ohrwurmes (Prolabia arachidis Yersin). Zeitschrift fiir Morpholo- gie und Okologie der Tiere 40: 158-180. https://doi.org/10.1007/ BF0042 1680 Houlbert CV (1900) Faune analytique illustrée des Orthoptéres de France. Deyrolle, Paris, 55 pp. [+ 12 pls.] ICZN — International Commission on Zoological Nomenclature (1999) International Code of Zoological Nomenclature. Fourth Edition. London: The International Trust for Zoological Nomenclature, c/o Natural History Museum. http://hdl.handle.net/1885/92062 Katoh K, Rozewicki J, Yamada KD (2019) MAFFT online service: multiple sequence alignment, interactive sequence choice and visu- alization. Briefings in bioinformatics 20(4): 1160-1166. https://dot. org/10.1093/bib/bbx 108 Kawano K (2006) Sexual dimorphism and the making of over- sized male characters in beetles (Coleoptera). Annals of the En- tomological Society of America 99(2): 327-341. https://doi. org/10.1603/0013-8746(2006)099[0327:SDATMO]2.0.CO;2 Kirby WF (1904) A synonymic catalogue of Orthoptera (Vol. I). Or- thoptera Euplexoptera, Cursoria, et Gressoria. (Forficulidae, Hemi- meridae, Blattidae, Mantidae, Phasmidae). British Museum (Natural History), London, 501 pp. Kirstova M, Kundrata R, Koéarek P (2020) Molecular phylogeny and classification of Chelidurella Verhoeff, stat. restit. (Dermap- tera: Forficulidae). Insect Systematics and Evolution, online 1-37. https://doi.org/10.1163/18763 12X-bjal0004 Klass KD (2001) The female abdomen of the viviparous earwig Hemi- merus vosseleri (Insecta: Dermaptera: Hemimeridae), with a dis- cussion of the postgenital abdomen of Insecta. Zoological Journal of the Linnean Society 131(3): 251-307. https://doi.org/10.1006/ zjls.2001.0246 247 Koéarek P (2004) Chelidura farkaci sp. n. (Dermaptera: Forficulidae) from China. Klapalekiana 40: 139-142. Lanfear R, Frandsen PB, Wright AM, Senfeld T, Calcott B (2016) PartitionFinder 2: new methods for selecting partitioned models of evolution for molecular and morphological phylogenetic analy- ses. Molecular Biology and Evolution 34(3): 772-773. https://doi. org/10.1093/molbev/msw260 Lapeira A, Pascual F (1980) Estudio preliminar de los dermapteros de la fauna ibérica. Trabajos Monograficos del Departamento de Zoologia de la Universidad de Granada, (n.s.) 3(4) [1977]: 53-99. Leraut P (2007) Insectos de Espafia y Europa. Lynx Editions, Bellaterra. 528 pp. [Spanish edition of Le guide entomologique. Delachaux-Ni- estlé, Paris]. Maccagno T (1933) I Dermatteri Italiani. Bollettino dei Musei di Zoo- logia ed Anatomia Comparata della R. Universita di Torino (ser. 3) 43(40): 241-296. Marquet M (1877) Notes pour servir a |’ Histoire Naturelle des insectes Orthopteres du Languedoc. Bulletin de la Société d’ Histoire Na- turelle de Toulouse 11: 137-159. Martorell Pefia M (1879) Catalogos sinonimicos de los insectos encon- trados en Catalufia, aumentados con los recientemente hallados por el Autor, en los diversos ordenes de los Coleopteros, Hemipteros, Hymenopteros, Ortopteros, Lepidopteros, Dipteros y Neuropteros. Tipogr. Sucesores N. Ramirez, Barcelona, 200 pp. Matzke D, Klass KD (2005) Reproductive biology and nymphal devel- opment in the basal earwig Jagalina papua (Insecta: Dermaptera: Pygidicranidae), with a comparison of brood care in Dermaptera and Embioptera. Entomologische Abhandlungen 62(2): 99-116. Miller MA, Pfeiffer W, Schwartz T (2010) Creating the CIPRES Sci- ence Gateway for inference of large phylogenetic trees. Proceedings of the Gateway Computing Environments Workshop (GCE), New Orleans, IEEE, 8 pp. https://doi.org/10.1109/GCE.2010.5676129 Mourier H (1986) Notes on the life history of Labia minor (L.) (Der- maptera), a potential predator of housefly eggs and larvae (Diptera, Musca domestica L.). Entomologiske Meddelelser 53(3): 143-148. Navas L (1921) Mis excursiones cientificas del verano de 1919. Memo- rias de la Real Academia de Ciencias y Artes de Barcelona 17[1920]: 143-169. Novellas F (1901) Ortopters de Catalunya. Catalech dels Ortopters ob- servats en aquesta regi0, fins el promer de janer de 1901. Butlleti de la Institucio Catalana d’ Historia Natural 1(4): e28. [37.] Ollason JG (1970) Studies on structural variation in the common Eu- ropean earwig (Forficula auricularia Linn). PhD Thesis, Durham University. http://etheses.dur.ac.uk/8617/ Pascual F (1988) 31 Isoptera, dermaptera, Phasmida y Embioptera [Pp. 415-443]. In: Barrientos JA (coord.) Bases para un curso practico de En- tomologia. Asociacion Espafiola de Entomologia, Salamanca, 754 pp. Plate I (1987) Italian and European Dermaptera of the museum of Flor- ence and Milan [Pp. 162—164]. In: Baccetti B (Ed.) Evolutionary biology of Orthopteroid Insects 157, E. Horwood, London, 1028 pp. Popham EJ (1968) Genera and species of Dermaptera (8) Eudohrninae, Neolobophorinae, Ancistrogastrinae, Diaperasticinae and Anechuri- nae (Forficulidae). The Entomologist 101: 196-201. Rambaut A, Drummond AJ, Xie D, Baele G, Suchard MA (2018) Tracer v.1.7. http://tree.bio.ed.ac.uk/software/tracer Ronquist F, Teslenko M, Van der Mark P, Ayres DL, Darling A, Hohna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MrBayes 3.2: dez.pensoft.net 248 efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61: 539-542. https://dot. org/10.1093/sysbio/sys029 Sakai S (1973) Dermapterorum Catalogus Praeliminaris. VII: A basic survey for numerical taxonomy pf the Forficulidae, the Dermaptera of the World. Special Bulletin, Daito Bunka University 6: 1-357. Sanchez-Vialas A, Garcia-Paris M, Ruiz JL, Recuero E (2020) Pat- terns of morphological diversification in giant Berberomeloe blis- ter beetles (Coleoptera: Meloidae) reveal an unexpected taxonomic diversity concordant with mtDNA phylogenetic structure. Zoologi- cal Journal of the Linnean Society 189(4): 1249-1312. https://dot. org/10.1093/zoolinnean/zlz164 Serville JGA (1831) Revue méthodique des Insectes de l’ordre des Or- thopteres. Annales de Sciences Naturales 22: 28-167. Serville JGA (1839) Histoire naturelle des insects — Orthopteres. Librai- rie encyclopédique de Roret, Paris, 777 pp. Shah AK (1984) The Biology of Forcipula trispinosa (Dohrn) (Der- maptera: Labiduridae). Records of the zoological survey of India. Miscellaneous publication. Occasional paper 53: 1-38. Simpson GB, Mayer DG (1990) Morphometric analysis of variation in Nala lividipes (Dufour) and Labidura truncata Kirby (Dermaptera: Labiduridae). Australian Journal of Entomology 29(4): 287-294. https://doi.org/10.1111/).1440-6055.1990.tb00365.x Srivastava GK (1970) Notes on a collection of Dermaptera from In- dia (Dermaptera). Eos, Revista Espafiola de Entomologia 45(1-4) [1969]: 319-333. Steinmann H (1981) The Dermaptera of the Universitets Zoologiske Museum, Kovenhavn. Folia Entomologica Hungarica 42: 173-192. Steinmann H (1989) World catalogue of Dermaptera. Series Entomo- logica, 43. Kluwer Academic Publishers, Dordrecht, Boston, Lon- don, 934 pp. dez.pensoft.net Pilar Jurado-Angulo et al.: Pyrenean species of Chelidura Steinmann H (1993) Dermaptera: Eudermaptera Il. Tierreich 108. Walter de Gruyter, Berlin, New York, 714 pp. https://dot. org/10.1515/9783 110872705 Swofford DL (2002) PAUP*: phylogenetic analysis using parsimony (* and other methods). Sinauer, Sunderland. Tomkins JL, Simmons LW (1996) Dimorphisms and _ fluctuating asymmetry in the forceps of male earwigs. Journal of Evolu- tionary Biology 9(6): 753-770. https://doi.org/10.1046/j.1420- 9101.1996.9060753.x Vancassel M (1984) Plasticity and adaptive radiation of dermapteran pa- rental behavior: results and perspectives [Pp. 51—79]. In: Rosenblatt JS, Beer C, Bunsen MC, Slater PJB (Eds) Advances in the Study of Behavior, 14. Academic Press, Orlando Fla., 237 pp. https://dot. org/10.1016/S0065-3454(08)60299-5 Vancassel M, Foraste M (1980) Le comportement parental des Der- mapteres. Reproduction Nutrition Développement 20(3B): 759- 770. https://doi.org/10.1051/rnd: 19800502 Wiley EO (1978). The evolutionary species concept reconsidered. Sys- tematic Zoology 27: 17-26. https://dot.org/10.2307/2412809 Wiley EO (1981). Phylogenetics. The theory and practice of phyloge- netic systematics. University of Kansas, Lawrence & John Wiley and sons, New York, 439 pp. Wiley EO, Mayden RL (2000) The evolutionary species concept. Q. D. Wheeler & R. Meier. Species Concept and Phylogenetic Theory: A Debate. Columbia University Press. New York, 70-92. Xambeu V (1903) Instinct de la maternité Chez le Chelidura dilatata, Lafrenaye, Orthoptere du groupe des Forficuliens. Le Naturaliste 25: 143-144. Xambeu V (1907) Notes explicatives du catalogue de la Faune des en- virons de Ria. Faune entomologique des Pyrénées-Orientales. Or- thoptéres. L’Echange, Revue Linneene, 275 (supplément): 204-208.