ZooKeys 217: 11–51, doi: 10.3897/zookeys.217.3136
Molecular phylogeny of the Trechus brucki group, with description of two new species from the Pyreneo-Cantabrian area (France, Spain) (Coleoptera, Carabidae, Trechinae)
Arnaud Faille 1,†, Charles Bourdeau 2,‡, Javier Fresneda 3,4,§
1 Zoologische Staatssammlung München, Münchhausenstraße 21, 81247 Munich, Germany
2 5 chemin Fournier-Haut, F–31320 Rebigue, France
3 Ca de Massa, 25526 Llesp – El Pont de Suert, Lleida, Spain
4  Museu de Ciències Naturals (Zoologia), Passeig Picasso s/n, 08003 Barcelona, Spain

Corresponding author: Arnaud Faille (faille@mnhn.fr)

Academic editor: Achille Casale

received 28 March 2012 | accepted 8 June 2012 | Published 28 August 2012

(C) 2012 Arnaud Faille. This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

For reference, use of the paginated PDF or printed version of this article is recommended.


A molecular phylogeny of the species from the Trechus brucki clade (previously Trechus uhagoni group)based on fragments of four mitochondrial genes and one nuclear gene is given. We describe Trechus (Trechus) bouilloni sp. n. from the western pre–Pyrenees: Sierras de Urbasa–Andía, Navarra, Spain. The species was collected in mesovoid shallow substratum (mss), a subterranean environment. Molecular as well as morphological evidences demonstrate that the new species belongs to the Trechus brucki clade. A narrow endemic species of high altitude in western French Pyrenees merged with Trechus brucki Fairmaire, 1862a, Trechus bruckoides sp. n., is described. A lectotype is designated for Trechus brucki and Trechus planiusculus Fairmaire, 1862b (junior synonym of Trechus brucki). The species group is redefined based on molecular and morphological characters, and renamed as the brucki group, as Trechus brucki was the first described species of the clade. A unique synapomorphy of the male genitalia, a characteristic secondary sclerotization of the sperm duct, which is shared by all the species of the brucki group sensu novo, is described and illustrated. The Trechus brucki group sensu novo is composed of Trechus beusti (Schaufuss, 1863), Trechus bouilloni sp. n., Trechus brucki, Trechus bruckoides sp. n., Trechus grenieri Pandellé, 1867, T. uhagoni uhagoni Crotch, 1869, T. uhagoni ruteri Colas, 1935 and Trechus pieltaini Jeannel, 1920. We discuss the taxonomy of the group and provide illustrations of structures showing the differences between the species, along with distribution data and biogeographical comments.


Carabidae, Trechini, Trechus brucki group , new species, molecular phylogeny, subterranean environment, Pyrenees, France, Spain


The genus Trechus (Coleoptera, Carabidae, Trechinae) includes more than 800 species, most of them in the Palearctic area (Moravec et al.2003; Lorenz 2005). This genus is known to contain many wingless short range endemic species (Jeannel 1927, Casale and Laneyrie 1982, Schmidt 2009) and is currently understood as polyphyletic (Faille et al. 2010a, 2011a).

Jeannel (1927) gathered seven species from the uhagoni group distributed from the French slope of the Pyrenees to the Cantabrian area: Trechus bonvouloiri Pandellé, 1867 (France: Hautes–Pyrénées), Trechus bordei Peyerimhoff, 1909 (France: Pyrénées–Atlantiques), Trechus brucki Fairmaire, 1862 (France: Hautes–Pyrénées, Pyrénées–Atlantiques), Trechus grenieri Pandellé, 1867 (France: Hautes–Pyrénées), Trechus navaricus Vuillefroy, 1867 (France: Pyrénées–Atlantiques), Trechus sharpi Jeannel, 1921 (Spain: Cantabria) and Trechus uhagoni Crotch, 1869 (Spain: Navarra). This group of species was also considered close to the group of Trechus angusticollis Kiesenwetter, 1850, a Pyreneo–Cantabrian group with nine species, all apterous, orophilous or troglobitic.

In this paper we describe a species collected by traps in a MSS (mesovoid shallow substratum, “Milieu Souterrain Superficiel” sensu Juberthie et al. 1980, Giachino and Vailati 2010) in the Sierras de Urbasa–Andía (Western pre–Pyrenees, Navarra, Spain) and a second orophilic species from the French Central Pyrenees. We study the phylogenetic relationships of the new species and provide a molecular phylogeny of the group, including all known species but four.

Historical background

Trechus uhagonii was described by Crotch in 1869 and dedicated to S. de Uhagon with whom he visited caves in the Alsasua area in June 1869. Trechus bruckii was first described under the name Trechus politus by Fairmaire (1862b). He renamed it one year later to bruckii because politus was already in use for an American species. The two names were corrected to uhagoni and brucki by subsequent authors, and recently renamed uhagonii and bruckii in catalogues (Lorenz 1998, 2005, Moravec et al. 2003, Queinnec and Ollivier 2011). As the names uhagonii and bruckii have not been used since their description, we choose to keep the prevailing usage of uhagoni and brucki in accordance with the article 33.3.1 of the International Code of Zoological Nomenclature on incorrect subsequent spellings (ICZN 1999).

In the Monographie des Trechinae, Jeannel (1927) erected the uhagoni group for the seven species of Trechus from the Pyreneo–Cantabrian area quoted above. Español (1970) described Trechus ortizi from a cave of Burgos province (Spain), and included it in the uhagoni group, close to Trechus bordei.

The uhagoni group sensu Jeannel (1927), although poorly defined morphologically, was enriched with 5 species by Casale and Laneyrie (1982) in their catalogue of species of world Trechinae: Trechus pecoudi Colas & Gaudin, 1935 (described first as a subspecies of Trechus brucki), Trechus ortizi, Trechus escalerae Abeille de Perrin, 1903 (considered by Jeannel (1927) to belong to the Trechus angusticollis group), Trechus enigmaticus Coiffait, 1971 and Trechus aubryi Coiffait, 1953. Trechus uhagoni was here considered as subspecies of Trechus grenieri. By describing Trechus baztanensis from a cave of Navarra, Dupré (1991) suggested that the peculiar genital morphology of Trechus bordei, Trechus navaricus, Trechus bonvouloiri and the new species should lead to their removal from the uhagoni group, and he created the bonvouloiri group for these species, opinion followed by Queinnec and Ollivier (2011). Toribio and Rodríguez (1997) added one species from Cantabria to the uhagoni group, Trechus carrilloi, a species collected in the MSS. Sciaky (1998) described Trechus jeannei from Cantabria, and included it in the uhagoni group, close to Trechus bordei and Trechus ortizi. Hernando (2002) described Trechus comasi from a cave of Navarra and suggested that it should be considered as sister species of Trechus brucki. Molecular and morphological evidence suggest that this species should be removed from the uhagoni group (Faille et al. 2010a, 2011a, Ortuño and Arribas 2010). Ortuño and Toribio (2005) described a new species of Trechus belonging to another group of species, indicating that 11 species belong to the uhagoni group in the Iberian Peninsula. Reboleira et al. (2010) considered that 10 species belong to this group in the Peninsula, without providing the list of taxa included.

Trechus brucki is an alpine species located at high altitude in the central and western Pyrenees, and it is until now not recorded from the Spanish slope of the chain (Serrano 2003). Colas and Gaudin described Trechus pecoudi in 1935 from the western Pyrenees (Pic d´Orhy) as a subspecies of Trechus brucki. Coiffait (1952) described 3 subspecies of Trechus brucki; Trechus brucki vandeli, Trechus brucki truilheti and Trechus brucki microthorax. The subspecies vandeli and truilheti were later related to Trechus pecoudi (Casale & Laneyrie, 1982) so that Trechus pecoudi counts three subspecies in the Catalogue of Palearctic Coleoptera (Moravec et al. 2003). Queinnec and Ollivier (2011) considered Trechus pecoudi as a subspecies of brucki restricted in the Anie and Orhy massifs, whereas the subspecies vandeli, described from Anie, was considered a synonym of Trechus brucki brucki together with the subspecies truilheti and microthorax.

Materials and methods Taxon sampling, Morphological study, DNA extraction and sequencing

Specimens were collected by hand or by means of pitfall traps containing water saturated in salt or propylene glycol, known to preserve DNA (Rubink et al. 2003, López and Oromí 2010) (Table 1). The protocol is detailed in Faille et al. (2010b):

Extractions of single specimens were non–destructive, using the DNeasy Tissue Kit (Qiagen GmbH, Hilden, Germany). After extraction, specimens were mounted on cards and genitalia stored in water–soluble dimethyl hydantoin formaldehyde resin (DMHF) on transparent cards, pinned beneath the specimen. Vouchers and DNA samples are kept in the collections of ZSM, IBE and MNHN.

We included examples of most species of the Trechus uhagoni group, with the exception of Trechus bruckoides sp. n., Trechus carrilloi and Trechus sharpi and some examples of Trechus of the angusticollis group sensu Jeannel (1927) and Casale and Laneyrie (1982) (Table 1). The tree was rooted with Aphaenops leschenaulti Bonvouloir, 1862, which is known to belong to a different group of Pyrenean Trechini (Jeannel 1927, Faille et al. 2010a).

We amplified fragments of four mitochondrial genes: 3’ end of cytochrome c oxidase subunit (cox1); a single fragment including the 3’ end of the large ribosomal unit (rrnL), the whole tRNA–Leu gene (trnL) and the 5’ end of the NADH dehydrogenase 1 (nad1);and one nuclear gene (internal fragment of the large ribosomal unit 28S rRNA, LSU) (see Table 2 for primers used). Sequences were assembled and edited using Sequencher TM 4.8 (Gene Codes, Inc., Ann Arbor, MI). Parts of the sequences for 14 of the species were taken from Faille et al. (2010a) and Faille et al. 2011a (Table 1).

New sequences have been deposited in the EMBL database with Accession Numbers HE817887–HE817940 (Table 1).

Table 1.

Sequenced specimens, with localities, collectors, codes and sequence accession numbers (unpublished sequences in bold).

sp locality collector code LSU cox1 rrnL trnL NAD1
Aphaenops Bonvouloir, 1862
Aphaenops leschenaulti Bonvouloir, 1861 Grotte de Castelmouly – Bagnères–de–Bigorre (France–65) C. Bourdeau, P. Déliot, A. Faille MNHN–AF1 GQ293593 HE817919 GQ293739 GQ293757 GQ293822
Trechus Clairville, 1806
Trechus grenieri Pandellé,  1867 Résurgence de la Hèche, Fréchet–Aure (France–65) J.P. Besson, C. Bourdeau, A. Faille ZSM–L13 HE817904 HE817920 HE817887 HE817887 HE817887
Trechus brucki Fairmaire,  1862 Pic du Gabizos, Arrens (France–65) C. Bourdeau ZSM–L329 HE817906 HE817921 HE817888 HE817888 HE817888
Trechus brucki Fairmaire,  1862 Pic du Gabizos, Arrens (France–65) C. Bourdeau ZSM–L329b HE817907 HE817889 HE817889 HE817889
Trechus brucki Fairmaire,  1862 Pic de Sesques, Laruns (France–64) C. Bourdeau ZSM–L446 HE817905 HE817922 HE817890 HE817890 HE817890
Trechus brucki Fairmaire,  1862 Pic de Gaziès, Laruns (France–64) C. Bourdeau ZSM–L190 HE817908 HE817923 HE817891 HE817891 HE817891
Trechus brucki Fairmaire,  1862 Caperan d´Anéou, Laruns (France–64) C. Bourdeau ZSM–L449 HE817909 HE817924 HE817892 HE817892 HE817892
Trechus uhagoni Crotch,  1869 Cueva de Orobe – Alsasúa (Spain–Navarra) C. Bourdeau, J. Fresneda ZSM–L161 HE817910 HE817925 HE817893 HE817893 HE817893
Trechus bouilloni Faille, Bourdeau & Fresneda, sp. n. Puerto de Lizarraga, Lizarraga (Spain–Navarra) C. Bourdeau, J. Fresneda ZSM_L201b HE817911 HE817926 HE817894 HE817894 HE817894
Trechus bouilloni Faille, Bourdeau & Fresneda, sp. n. Puerto de Lizarraga, Lizarraga (Spain–Navarra) C. Bourdeau, J. Fresneda ZSM_L201t HE817927 HE817895 HE817895 HE817895
Trechus beusti (Schaufuss 1863) Cueva de San Adrián, Zegama (Spain–Guipúzcoa) C. Bourdeau, J. Fresneda ZSM–L199 HE817912 HE817928 HE817896 HE817896 HE817896
Trechus pieltaini Jeannel,  1920 Cueva de Mairuelegorreta, Gorbea (Spain–Álava) C. Bourdeau ZSM–L395 HE817913 HE817929 HE817897 HE817897 HE817897
Trechus navaricus (Vuillefroy, 1869) Grotte de Sare – Sare (France–64) C. Bourdeau MNHN–AF103 GQ293603 GQ293687 FR729578 FR729578 FR729578
Trechus bordei Peyerimhoff,  1909 Grotte d´Ayssaguer – Larrau (France–64) C. Bourdeau, P. Déliot, A. Faille MNHN–TBA HE817914 HE817930 HE817898 HE817898 HE817898
Trechus bonvouloiri Pandellé, 1867 Pic de Montaigu – Baudéan (France – 65) C. Bourdeau ZSM–L218 HE817915 HE817931 HE817899 HE817899 HE817899
Trechus abeillei Pandellé,  1872 Cirque d´Anglade Couflens (France–09) C. Vanderbergh ZSM–L15 HE817916 HE817932 HE817900 HE817900 HE817900
Trechus distinctus Fairmaire & Laboulbène, 1854 Col Sobe Ariel – Laruns (France–64) C. Bourdeau ZSM–L216 HE817917 HE817933 HE817901 HE817901 HE817901
Trechus aubryi Coiffait,  1953 Cirque d´Anglade Couflens (France–09) B. Junger ZSM–L370 HE817934 HE817902 HE817902 HE817902
Trechus jeannei Sciaky,  1998 Bosque de Saja, Saja (Spain–Cantabria) C. Bourdeau ZSM–L516 HE817918 HE817903 HE817903 HE817903
Trechus saxicola Putzeys,  1870 Braña Caballo – Piedrafita (Spain–León) C. Bourdeau, P. Déliot, A. Faille MNHN–AF100 GQ293614 HE817935 FR729577 FR729577 FR729577
Trechus escalerae Abeille de Perrin, 1903 Cueva de Porro Covañona – Covadonga (Spain–Asturias) J.M. Salgado MNHN–AF104 GQ293612 FR733912 GQ293731 GQ293793 GQ293839
Trechus ceballosi Mateu,  1953 Aven de Licie Etsaut – Lanne–en–Barétous (France–64) C. Bourdeau, A. Faille MNHN–AF128 GQ293610 FR733914 GQ293728 GQ293791 GQ293850
Trechus distigma Kiesenwetter, 1851 Aven de Nabails – Arthez d’Asson (France–64) C. Bourdeau, P. Déliot, A. Faille MNHN–AF94 GQ293611 HE817936 FR729575 FR729575 FR729575
Trechus barnevillei Pandellé,  1867 Cueva del Pis – Penilla, Santiurde de Toranzo (Spain–Cantabria) C. Bourdeau, P. Déliot, A. Faille MNHN–AF97 GQ293607 GQ293680 GQ293727 GQ293783 GQ293848
Trechus obtusus Erichson,  1837 Estrada de Nicho (Portugal–Madeira) A. Arraiol IBE–AF2 FR733997 HE817937 FR729579 FR729579 FR729579
Trechus quadristriatus (Schrank, 1781) Collau de la Plana del Turbón – Egea (Spain– Huesca) P. Déliot, A. Faille, J. Fresneda MNHN–AF96 GQ293619 FR733908 GQ293743 GQ293745 GQ293841
Trechus fulvus Dejean,  1831 Cueva del Pis – Penilla, Santiurde de Toranzo (Spain–Cantabria) C. Bourdeau, P. Déliot, A. Faille MNHN–AF98 GQ293613 HE817938 GQ293729
Trechus martinezi Jeannel,  1927 Cova de les Meravelles – Cocentaina (Spain– Alicante) C. Andújar, P. Arribas, A. Faille IBE–AF1 FR733996 HE817939 FR729576 FR729576 FR729576
Trechus schaufussi ssp. comasi Hernando, 2002 Cueva Basaula – Barindano (Spain–Navarra) J. Fresneda MNHN–AF127 GQ293617 HE817940 FR729580 FR729580 FR729580
Apoduvalius Jeannel, 1953
Apoduvalius alberichae Español, 1971 Cova de Agudir – Cardano de abajo – Palencia (Spain–Asturias) J.M. Salgado MNHN–AF105 GQ293618 GQ293632 GQ293732 GQ293794 GQ293840
Apoduvalius anseriformis Salgado et Peláez, 2004 Cueva de Entrecuevas – Caravia Alta (Spain– Palencia) A. Cieslak, A. Faille, J. Fresneda, I. Ribera, J.M. Salgado MNCN–AF2 FR733999 FR733916 FR729582 FR729582 FR729582
Table 2.

Primers used in the study. F, forward; R, reverse.

Gene Name Sense Sequence Reference
cox1 Jerry (M202) F CAACATTTATTTTGATTTTTTGG Simon et al. 1994
Chy F T(A/T)GTAGCCCA(T/C)TTTCATTA(T/C)GT Ribera et al. 2010
Tom R AC(A/G)TAATGAAA(A/G)TGGGCTAC(T/A)A Ribera et al. 2010
rrnLnad1 16saR (M14) F CGCCTGTTTA(A/T)CAAAAACAT Simon et al. 1994
Phylogenetic analyses

We aligned the sequences using the MAFFT online v.6 and the Q–INS–i algorithm (Katoh and Toh 2008), a progressive pair–wise method with secondary refinement. We used Maximum Likelihood as implemented in the on–line version of RAxML (which includes an estimation of bootstrap node support, Stamatakis et al. 2008), using GTR+G as the evolutionary model and three partitions corresponding to the cox1, rrnL+trnL+nad1 and LSU fragments.

The aedeagus and genital duct were extracted and included in a drop of Canada balsam or dimethyl hydantoin formaldehyde resin (DMHF) on a transparent slide. Preparations were mounted below the specimen, on the same pin. Pictures were taken with microscopes Olympus ch and Olympus szx16, coupled with a camera Olympus c5060wz. Serial pictures were combined using the CombineZP software, and finally processed using Adobe Photoshop CS.

Institutional codes and abbreviations used in the taxonomic treatment and private collectors

IBE Institute of Evolutionary Biology (CSIC-UPF), Barcelona (Spain).

MNCN Museo Nacional de Ciencias Naturales (CSIC), Madrid (Spain)

MNHN Muséum National d´Histoire Naturelle, Paris (France).

MZB Museu de Ciències Naturals (Zoologia), Barcelona (Spain).

ZSM Zoologische Staatssammlung, München (Germany).

MFN Museum für Naturkunde, Berlin (Germany).

CAF coll. A. Faille (Paris, France).

CCB coll. C. Bourdeau (Rebigue, France).

CJF coll. J. Fresneda (Llesp, Spain).

CMT coll. M. Toribio (Madrid, Spain).

LE Length of elytra.

LP Length of pronotum.

WE Width of elytra.

WH Width of head.

WP Width of pronotum.

WPB Width of pronotal base.

Trechus bouilloni Faille, Bourdeau & Fresneda, sp. n.



Figs 1, 8, 15, 16,29
Type locality.

Spain, Navarra, Sierra de Urbasa–Andía, Lizarraga, Puerto de Lizarraga, UTM (WGS 84): 30 T, X: 580, Y: 4746, Z: 900 m.

Type series.

Holotype (MNHN): 1 ♂, Spain, Navarra, Sierra de Urbasa–Andía, Lizarraga, Puerto de Lizarraga, MSS, trap: 1–5–1980/15–8–1980, Bourdeau and Fresneda leg., voucher number ZSM–L201, MNHN]. DNA aliquotes preserved in the DNA and tissue collections of the ZSM, MNHN and IBE; Genitalia dissected and mounted in a separate label pinned with the specimen. Paratypes: 52 ♂♂, 62 ♀♀, same label data as holotype (MNCN, MNHN, MZB, ZSM, CCB, CJF, CAF, CMT).


Large size (ca 5 mm) and round shape (Fig. 1). Median lobe of aedeagus slender, in lateral view (Fig. 15) the basal third curved, the central part straight and the apex with a curved hook assymetrical in dorsal view (Fig. 8). Inner sac of aedeagus (=endophallus) with an elongate and well-sclerotized piece, forming a gut and armed with internal scales. Characteristic secondary sclerotization of the sperm duct (Fig. 15: CP2) forming a kind of second copulatory piece outside base of the median lobe.

Description of the holotype.

Habitus as in Fig. 1.Elongated, round–sided. Body surface with a very thin, hardly visible, dense microreticulation, with more distinguishable meshes on the head.

Colour. Dorsal surface dark brown, moderately shiny. Antennae, palpi and legs light brown.

Chetotaxy. Surface of elytra glabrous with the exception of a periscutellar seta, two discal setae on the third stria, four humeral setae, four setae along lateral margin and two preapical setae. Marginal setae of pronotum present, the anterior ones located before the first third of the length. Ventral pubescence limited to one seta on each half sternite.

Head. Eyes reduced, flat; ommatidia well defined; maximum diameter of about eight ommatidia, temples approximately twice the length of eyes, strongly wrinkled to the neck. Frontal furrows deeply impressed. Antennae moderately long, five antennomeres extend beyond the pronotal base. Antennomere III distinctly longer than antennomeres II and IV, which are similar in length.

Pronotum. Proportions (M–F): WP/LP = 1.3–1.28, WP/WPB = 1.3–1.3, WP/WH = 1.38–1.3, WE/WP = 1.57–1.53. Transverse, with lateral margins finely bordered; wider in anterior part, narrower than elytra; posterior part much narrower than base of elytra. One seta in the marginal gutter at about a third of pronotum length, another one close to hind angle. Sides evenly rounded and straight just between hind angles and insertions of posterior setae. Hind angles well developed, salient.

Elytra. Proportions (M–F): WE/LE = 0.65–0.69. Oval, broadest almost at mid–length; surface moderately convex, flattened on disc. Shoulders distinct but rounded. Striae very finely punctuated, sixth inner striae deeply impressed on disc, but reduced at apex and base; seventh striae shallower, but distinct, the eighth reduced to the posterior quarter of elytra. Apical striola strongly impressed continuing the fifth stria.

Hind wings. Very reduced, not functional.

Male genitalia. Median lobe of aedeagus slender, in lateral view (Fig. 15) the basal third curved, the central part straight and the apex showing a curved hook; assymetrical in dorsal view (Fig. 8). Parameres slender, each with 4 to 6 setae at apex. Internal sac of aedeagus with an elongate well-sclerotized piece, forming a symmetrical gut and armed with internal scales (Fig. 16). Characteristic secondary sclerotization of the sperm duct forming a kind of second copulatory piece out of the base of the median lobe (Fig. 15: CP2).

Female genitalia.

Internal genitalia membranous. Gonocoxites unguiform, with 4 to 5 large setae, and 2 small near apex. Gonosubcoxites with 2 to 3 large setae near the internal edge. Laterotergite IX with 12 setae at the basal margin, and 4 to 6 scattered (Fig. 29).


Mean length (5 exemplars): 5.25 mm (male), 4.56 mm (female).


The new species is dedicated to Michel Bouillon, Pyrenean speleologist, who was the first to discover the existence of cave beetles in MSS.


Trechus bouilloni sp. n. is a representative of the Trechus brucki group sensu novo as defined in the present paper. It shares with Trechus grenieri, Trechus uhagoni, Trechus beusti, and Trechus pieltaini the same kind of aedeagus morphology, especially the apex with a curved hook in lateral view, and an internal sac showing two sclerotized parts, the internal copulatory piece and another triangular piece forming a kind of second copulatory piece (CP2, Figs 1724), also existing in Trechus brucki and Trechus bruckoides sp. n. (Figs 25–28). Similar secondary sclerotized structures of endophallus are known in some groups of insects including Coleoptera, and described as a “sperm pump” (Beutel and Leschen 2005, Hünefeld and Beutel 2005, Jäch and Delgado 2010). In the Trechus brucki group, however, the structure is too rudimentary to play the same role in sperm transfer, and its function –if any– remains obscure. Although never observed before, this structure is also present in the others species of the group and is the main synapomorphy of the clade. The lack of this internal sclerotized structure in Trechus carrilloi and Trechus sharpi (Figs 30–32) casts doubt on their affinities.

Distribution and ecology.

Trechus bouilloni sp. n. is only known from the type locality, the MSS of Lizarraga pass (Navarra, Spain) (Fig. 36). The type locality is a MSS located on a northern slope at the eastern extremity of the Sierra de Andía–Urbasa, close to the Lizarraga pass.

Trechus were collected by means of traps in a zone of scree (altitude: 900 m) extending from east to west at the feet of cliffs of Albian limestone lining the northern slope of the plateau of the Sierra de Andía–Urbasa. This scree slope consists of a mass of fallen rocks resulting from the erosion of calcareous cliffs and constitutes a steeply sloped (45°) MSS, filling one of the numerous gullies of a beech forest covering the entire northern side of the plateau lining the southward depression of the Río Arakil (Sakana valley).

On this unstable ground, beeches are replaced by grassy and mossy vegetation dotted with shrubs. The layer of humus is irregular and very thin and only partly covers the blocks of white, angular, medium–sized limestone, rarely exceeding the size of 1 dm³.

The traps were placed 50 centimeters deep in a “C–type” horizon (sensu Juberthie et al. 1981), constituted mainly by stones of 5 cm³, not sealed by the ground and not welded, leaving numerous spaces between them and forming a layer several meters thick above the compact rock.

The other Coleoptera collected with Trechus bouilloni sp. n. were Leiodidae, Cholevinae: Catops subfuscus Kellner, 1846, Sciodrepoides watsoni (Spence, 1813) (Catopini) and Bathysciola sp. (Leptodirini).

Some specimens of Trechus bouilloni sp. n. were parasitized by an undetermined Ascomycete.

Trechus bouilloni sp. n. was not found in caves of the area north of Larraona (cueva de los Cristinos, cuevas de Erbeltz, Txintxoleze, Noriturri, Akuandi, del Queso, Iniriturri, Arleze, Laminatitur), suggesting that it is strictly located in MSS (CB personal observation).

Trechus brucki Fairmaire, 1862


Figs 7, 14, 27, 28, 33
Type locality.

«Eaux–Bonnes, M. vom Bruck» (Fairmaire, 1862b). France, Pyrénées–Atlantiques.

Type series.

Lectotype (MNHN), present designation: 1 ♂, labelled: «oblongulus Bonnes» [white rectangular label (ms, Fairmaire)], «Bruckii» [white rectangular label (ms, Fairmaire)], «MUSEUM PARIS Collection Léon Fairmaire 1906» [white rectangular label (printed)], «TYPE» [red rectangular label (printed)], «Lectotypus / Trechus bruckii Fairmaire / Faille, Bourdeau & / Fresneda des. 2012” [red rectangular label (printed)], genitalia dissected and mounted in a separate label pinned with the specimen. Paralectotype (MNHN): 1 ♀, same label data and pin as lectotype except “Paralectotypus / Trechus bruckii Fairmaire / Faille, Bourdeau & / Fresneda des. 2012” [red rectangular label (printed)].

Type series of Trechus planiusculus Fairmaire, 1862.

Lectotype (MNHN), present designation: 1 ♀ (red dot), labelled: “oblongus” [white rectangular label (ms, Fairmaire)], “planiusculus” [white rectangular label (ms, Fairmaire)], “Bruckii” [white rectangular label (ms, Fairmaire)], “2203” [white rectangular label (ms, Fairmaire)], “MUSEUM PARIS Collection Léon Fairmaire 1906” [white rectangular label (printed)], “TYPE” [red rectangular label (printed)], “Lectotypus / Trechus planiusculus Fairmaire / Faille, Bourdeau & / Fresneda des. 2012” [red rectangular label (printed)]. Paralectotypes (MNHN): 1 ♀, same label data and pin as lectotype except «Paralectotypus / Trechus planiusculus Frm / Faille, Bourdeau & / Fresneda des. 2012” [red rectangular label (printed)]; 1 ♂, “H Pyrenees 1856 M. Pandellé” [white rectangular label (printed)], “Bruckii” [white rectangular label (ms, Fairmaire)], “COTYPE” [white and red rectangular label (printed)], “R. Jeannel Brucki Fr” [white rectangular label (ms, Jeannel)], “MUSEUM PARIS coll. R. JEANNEL 1931” [white rectangular label (printed)], “Paralectotypus / Trechus planiusculus Frm / Faille, Bourdeau & / Fresneda des. 2012” [red rectangular label (printed)], genitalia dissected and mounted in a separate label pinned with the specimen.

Non Type material.

1 ♀ (MNHN) labelled: “planiusculus” [white rectangular label (ms, Fairmaire ?)], “Bruckii” [white rectangular label (ms, Fairmaire)], “MUSEUM PARIS Collection Léon Fairmaire 1906” [white rectangular label (printed)], “R. Jeannel Brucki Fr” [white rectangular label (ms, Jeannel)]. We do not consider this specimen as a syntype of Trechus planiusculus as it is not labeled « oblongus » as the specimen of the type series, suggesting that the specimen arrived in the Fairmaire collection after the description of planiusculus. A second female specimen (MNHN) labelled: “oblongus Arrens” [white rectangular label (ms, Fairmaire)], “TYPE” [white and red rectangular label (printed)],  “MUSEUM PARIS Collection Léon Fairmaire 1906” [white rectangular label (printed)]. This specimen could be the reference specimen of Trechus oblongus Schaum, 1862. Reference of the name comes from Schaum (1862: addenda, p. 119): “P. 14 col. 2 Trechus oblongus Schaum;” only the name is mentioned, without any description, number of exemplars studied or locality. It should then be considered as nomen nudum. Jeannel (1927) indicates that Trechus oblongus is a synonym of Trechus brucki with type locality: “Pyrén. occ.” We were unable to find the specimen or reference where Jeannel found the type locality.

Taxonomic comments

The study of specimens of Trechus brucki pecoudi from Orhy and of numerous exemplars of Trechus brucki, including types of the previously described subspecies of Trechus brucki, demonstrated that none of the characters quoted either by Colas and Gaudin (1935) or by Queinnec and Ollivier (2011) are constant. We consider then the subspecies pecoudi as synonymous of Trechus brucki: Trechus brucki brucki Fairmaire, 1862 = Trechus brucki pecoudi Colas & Gaudin, 1935, syn. n.

Trechus politus and Trechus planiusculus were described by Fairmaire in the volume of the Annales de la Société Entomologique de France of 1861 published in 1862 (Fairmaire, 1862b). As the name Trechus politus was already used for an American species (today Trechisibus politus Brullé, 1842), Fairmaire (1862a) changed the name of this species to Trechus bruckii. Trechus planiusculus was considered synonymous with Trechus brucki by Jeannel (1927), and, moreover, the name planiusculus was preoccupied as it was used by Costa (1858) in a work on Italian fauna. In his works on Trechini, Jeannel (1927, 1941) illustrated the genitalia of a male from the Ossau Valley. Recently, in a revision of the french fauna of Carabidae, Queinnec and Ollivier (2011) suggested that the drawing of Jeannel (1927, 1941) was incorrect, and that the male genitalia of Trechus brucki has a homogeneous shape throughout the distribution area. By examining the types of Fairmaire, we noticed that the drawing of Jeannel (1927, 1941) does not actually match with Trechus brucki. However, by studying specimens from Ossau Valley we found that the drawing of Jeannel actually corresponds to another undescribed species, very narrowly located in the area of Pic de Montagnon (Bielle–Pyrénées Atlantiques). Here we describe this new species as Trechus bruckoides sp. n.

Trechus bruckoides Faille, Bourdeau & Fresneda, sp. n.



Figs 6, 13, 25, 26
Type locality.

France, Pyrénées Atlantiques, Ossau, Sède de Pan UTM (WGS 84): 30 T, X:704, Y:4768.

Type series.

Holotype (MNHN): 1 ♂, France, Pyrénées Atlantiques, Ossau, Sède de Pan, labelled: «Ossau, Sède–Pan» [white rectangular label (printed)], «MUSEUM PARIS coll. R. JEANNEL 1931» [white rectangular label (printed)], «R. Jeannel Brucki Fr.» [white rectangular label (ms, Jeannel)], «Holotypus / Trechus bruckoides sp. n. / Faille, Bourdeau & / Fresneda det. 2012” [red rectangular label (printed)], genitalia dissected and mounted in a separate label pinned with the specimen. Paratypes: 1 ♂, “Pic Montagnoü (v. d´Ossau) Mascaraux” [white rectangular label (ms)], “MUSEUM PARIS 1932 coll. Sainte–Claire Deville” [white rectangular label (printed)], “angusticollis Kiesw.” [white rectangular label (ms)] (MNHN); 1 ♂, “Pic Massibe B. PYR. 1938” [white rectangular label (ms)], “Trechus Brucki” [white rectangular label (ms)], “Collection H. Coiffait” [white rectangular label (printed)] (MNHN); 1 ♂, “Bielle/ B. Pyr.” “Trechus brucki/det. Tedeschi” “coll. Tedeschi/ZSM 2009” (ZSM); Pic Montagnon, 15–VII–1979, Bourdeau leg., 6 ♂♂ and 1 ♀ (CAF, CCB, CJF); Sède de Pan, Bielle, VII–1995, Bourdeau leg., 1 ♂ (CCB); Sède de Pan, Bielle, 2–VIII–1980, Bourdeau leg., 3 ♂♂ (CCB); Sède de Pan, Bielle, 10–VII–1981, Bourdeau leg., 1 ♂ and 2 ♀♀ (CCB). All the paratypes with the label “Paratypus / Trechus bruckoides sp. n. / Faille, Bourdeau & / Fresneda det. 2012” [red rectangular label (printed)].

Supplementary specimen studied.

1 ♀, «Pied du pic Lauriolle près Bielle Bas. Pyr. 29.6.37», coll. Bonnaire (MNHN). Sède de Pan, Mascaraux, 2 exx. (coll. Nègre, MNHN). Pic Montagnon: 4 exx. Sède de Pan: 3 exx (MNHN). Sède de Pan, 23–6–1943, 1 ♂, 2 ♀♀ (MNHN, coll. Coiffait). Pic Massibe: VII–1941, 1 ♀ (MNHN, coll.Coiffait).


Large size (ca 4 mm) and round shape (Fig. 6). Median lobe of aedeagus slender, subparallel and decreasing in width from the apical tenth to the apex, which is softly curved in lateral view (Fig. 25), nearly symmetrical and with apex regularly rounded in dorsal view (Fig. 13). Endophallus with an elongate and well-sclerotized piece, forming a twisted gut. Characteristic secondary sclerotization of the sperm duct (Fig 25: CP2) present. External appearance very close to Trechus brucki.

Description of the holotype.

Habitus as in Fig. 6.Elongated, round–sided. Body surface with a very thin, hardly visible, dense microreticulation, no more distinguishable meshes on the head.

Colour. Dorsal surface dark brown, moderately shiny. Antennae, palpi and legs light brown.

Chetotaxy. Surface of elytra glabrous with the exception of a periscutellar seta, two discal setae on the third stria, four humeral setae, four setae along lateral margin and two preapical setae. Marginal setae of pronotum present, the anterior ones located at the first anterior third of the length.

Head. Eyes flat, well–developed, temples smaller than the length of eyes, strongly wrinkled to the neck. Frontal furrows moderately deep. Antennae short (2–2.3mm) and thick.

Pronotum. Proportions (M): WP/LP = 1.3, WP/WPB = 1.35, WP/WH = 1.34, WE/WP = 1.63. Transverse, with lateral margins bordered, wider in anterior part, much less wide than elytra. Posterior part much narrower than base of elytra. One seta in the marginal gutter at about a third of pronotum length, another one just before hind angle. Sides evenly rounded and straight just between hind angles and insertions of posterior setae. Hind angles well developed, right.

Elytra. Proportions (M): WE/LE = 0.64. Subrectangular, broadest after the mid–length; surface moderately convex, flattened on disc. Shoulders distinct but rounded. Striae almost impunctuate, sixth inner discal striae distinct, but reduced at apex and base, especially in callus area; seventh striae shallower, nearly indistinct, the eighth only distinct close to apex of elytra. Apical striola well impressed continuing the fifth stria.

Hind wings. Very reduced, not functional.

Male genitalia. Median lobe of aedeagus slender, in lateral view (Fig 25) basal third curved, central part straight, parallel and elongated towards apex. Nearly symmetrical in dorsal view (Fig 13). Parameres slender, each with 4 setae at tip. Inner sac of aedeagus armed with scales with an elongate well sclerotized piece, forming a twisted gut (Fig 26). Characteristic secondary sclerotization of the sperm duct forming a kind of second copulatory piece out of the base of the median lobe (Fig 25: CP2).

Female genitalia.

Not examined.


Mean length (4 exemplars): 4.78 mm (male).


The specific epithet refers to Trechus brucki, species with which the new species was merged.


Trechus brucki and Trechus bruckoides sp. n. are externally very similar but strong differences isolate the two taxa especially in shape of male genitalia (Figs 25, 27). The aedeagus shape of Trechus bruckoides sp. n. is exactly as indicated in Jeannel (1927, 1941) for Trechus brucki.

Distribution and ecology.

Trechus bruckoides sp. n. is only known from the calcareous plateau of Esturou located at 1860 m, north of Montagnon peak (1973 m) and Mailh Massibé (1973 m), at the northern extremity of the massifs separating Aspe and Ossau valleys (Fig. 37). South of this area (Sesques and Gaziès peaks (2600 m)), it is replaced by Trechus brucki which occurs together with Trechus distinctus. During Pleistocene glacial cycles, this plateau was covered by a névé which shaped an area of sinks of nivo–karstic origin (Auly 2008). After winter, snow remains in these sinkholes (July–August) and allows the preservation of a nivicolous fauna, which is unusual at these medium altitudes. Trechus bruckoides sp. n. lives exclusively in the masses of fallen rocks of sinkholes and follows the withdrawal of the snow. When the snow thaws it likely seeks refuge underground.

This mid altitude nivicolous environment could have led to isolation of populations of the species from southern glaciated areas and glacial tongues of the northern slope of Ossau glacier and led to the differentiation of this population of cryophilic and highly hygrophilic Trechus. Such a hypothesis could also explain the presence of the hypogean Trechini Aphaenops bessoni Cabidoche, 1962, endemic to this karstic plateau (pits of Col d’Aran), and closely related to Aphaenops loubensi Jeannel, 1953, an endemic species of the Pierre Saint Martin massif, western to the Aspe Valley. Some other endemic nivicolous Carabidae with morphologically distinct populations occur in the area, like Carabus (Iniopachus) pyrenaeus Audinet–Serville, 1821 (the population of Sède de Pan was first described as a distinct subspecies, Carabus pyrenaeus cephalicus Csiki, 1927), Nebria lafresnayei Audinet–Serville, 1821, Pterostichus (Cryobius) amoenus mascarauxianus Pupier, 2008, Pterostichus (Lianoe) nadari mascarauxi Jeannel, 1928 and Pterostichus (Lianoe) dufourii (Dejean 1828). The peculiarities of this fauna suggest that this restricted area is an important center of diversification.


The molecular phylogeny (Fig. 34) suggests a well–supported clade gathering the following species:

Trechus beusti (Fig. 4), Trechus bouilloni sp. n.(Fig. 1), Trechus grenieri (Fig. 2), Trechus brucki (Fig. 7), Trechus pieltaini (Fig. 5) and Trechus uhagoni (Fig. 3). This result is in accordance with morphology: all the species of the clade share the aedeagal median lobe long and strongly curved just behind basal bulb, with terminal lamella well–developed. Moreover, the clade is supported by a strong synapomorphy: all the species share a strongly sclerotized part of the sperm duct, forming a second copulatory piece (Figs 15, 17, 19, 21, 23, 27: CP2). This synapomorphy is also present in Trechus bruckoides sp. n.(Fig. 25). Consequently, molecular and morphological results allow us to define the Trechus brucki group sensu novo: Trechus beusti, Trechus bouilloni sp. n., Trechus brucki, Trechus bruckoides sp. n., Trechus grenieri, Trechus uhagoni and Trechus pieltaini.

Two species, Trechus pieltaini and Trechus beusti, were included by Jeannel (1927) in the angusticollis group. Trechus carrilloi and Trechus sharpi are provisionally not included in the group because of the absence of sclerotization of the sperm duct (CP2). Moreover, concerning Trechus carrilloi, the apical hook is not a synapomorphy, as several other Trechus groups have this kind of hook (i.e. Trechus aubryi or the Tibetan species Trechus bastropi Schmidt, 2009 and Trechus damchungensis Deuve, 1997). As expected by Dupré (1991) for some of the species that he included in a “bonvouloiri group”, the following regional species previously considered part of the brucki group are clearly excluded here: Trechus escalerai, Trechus navaricus, Trechus bordei, Trechus jeannei, Trechus bonvouloiri. Although not included in our analyses, we also exclude of the group Trechus ortizi, Trechus baztanensis and Trechus enigmaticus, as those species were put into the uhagoni group because of close morphological affinities with Trechus bordei and Trechus navaricus (Español 1970, Coiffait 1971, Dupré 1991). Trechus jeannei, which was said to be close to Trechus bordei (Sciaky 1998), does not belong to the Trechus brucki clade and is not clearly related with the Trechus bordei clade. A study including more Iberian species should clarify its phylogenetic affinities. Trechus aubryi is excluded from the Trechus brucki clade and shares strong affinities with Trechus distinctus.

Our molecular results as well as genital morphology, in dorsal and lateral view and shape of copulatory piece (Figs 10, 19, 20), suggest that Trechus uhagoni could be considered a distinct species from Trechus grenieri. Morphological differences between the two species are the following:

Trechus grenieri: aedeagus in dorsal view (Fig. 9) with subparallel sides, round apex with a short triangular tip; in lateral view (Fig. 17) basal third strongly rounded, median lobe slightly angular in the middle; apical hook with a thin tip. The copulatory piece is an asymmetrical gut slightly tapering and filled with a densely scaly area (Fig. 18).

Trechus uhagoni: aedeagus in dorsal view (Fig. 10) with the left side narrowed or sinuate from the middle to apex, the left side of apical quarter deeply narrowed, forming a long triangular tip; in lateral view (Fig. 19) only the basal quarter rounded, median lobe without dorsal angle in the middle, short with apical hook with massive tip. The copulatory piece is similar to the one of grenieri but the gut is parallel and shortened in its apical part (Fig. 20).

With 7 subspecies recognized in the last catalogues (Moravec et al 2003, Queinnec and Ollivier 2011), Trechus grenieri currently lives in humid forests (1000 m) from Espinal to Iraty (ssp. ruteri), then from Gave de Pau river (north side of Pic de Montaigu) to the Neste d’Aure valley (ssp. grenieri). From Aure valley to the Salat, it is replaced by the subspecies despaxi, which crosses the Garonne river near Saint Béat (Haute Garonne). An isolated subspecies (prepyrenaeus) was described by Coiffait in the high Arize valley (Andronne and Bosc forests around 1000 m) (Coiffait 1974). Along the axial ridge, Trechus grenieri lives above 1500 m from high Garonne valley (ssp. bepmalei) to Mont Valier (ssp. aulaensis Aubry, 1981). Study of numerous specimens suggests that Trechus uhagoni and all the specimens of the subspecies ruteri are morphologically close, especially in the shape of the male genitalia, and should be considered as a distinct species. Moreover, we studied specimens from various localities of the Pyrenean range (see Distribution) and established that they share some morphological characters (color pale, brown, pronotum transverse with lateral margin regularly rounded, elytral striae superficial, weakly impressed) that justify keeping the status of ruteri as a subspecies of Trechus uhagoni. The subspecies ruteri should then be considered as belonging to uhagoni, so that grenieri is restricted to the area between Gave de Pau and Ariège valley, northern slope of Pyrenees. Trechus uhagoni ruteri n. comb. could be distinguished from Trechus uhagoni uhagoni by its color, usually paler brown, the pronotum transverse with lateral margin regularly rounded and the elytral striae superficial, weakly impressed. It is restricted to the western Pyrenees.

The study of specimens from the whole range of Trechus grenieri including all the subspecies, most of the types and material from intermediate localities (see distribution) leads us to conclude that the characters used to discriminate the subspecies (size, eyes size, shape of elytra and pronotum) are inconstant and overlapping between populations. The shape of the male genitalia is similar for all the populations between Gave de Pau and Ariège valley, including the one (ssp aulaensis) which was said to be different (Queinnec and Ollivier 2011). We consider then Trechus grenieri as a single species without any valid subspecies: Trechus grenieri grenieri Pandellé, 1867 = Trechus grenieri bepmalei Jeannel, 1921 = Trechus grenieri despaxi Jeannel, 1922 = Trechus grenieri aulaensis Aubry, 1981 = Trechus grenieri prepyrenaeus Coiffait, 1974, syn.n.

Trechus beusti was described by Schaufuss (1863). The type series is located in the Schaufuss collection in the Museum für Naturkunde, Berlin (M. Jaeger pers. com.). Trechus pieltaini was described by Jeannel (1920) from a cave of the Basque country, Cueva de Mairruelegorreta. Bolívar y Pieltain and Jeannel (1921) suggested that the peculiar morphology of the aedeagus of these two species indicates clear affinities with Trechus uhagoni and Trechus grenieri. Surprisingly, Jeannel (1927) in his Monographie des Trechinae considered that these two species belong to another group of species, the Trechus angusticollis group. This opinion was followed by subsequent authors (Español 1965, Casale and Laneyrie 1982, Ortuño and Marcos 2003). However, and in accordance with the morphology of the median lobe of the aedeagus, molecular results support Bolívar y Pieltain and Jeannel’s (1921) point of view and confirm the close affinities between Trechus beusti, Trechus pieltaini and thespecies of the Trechus brucki clade. Differences between the two species are weak: the apical part of the aedeagus is longer in Trechus pieltaini (Fig. 12, 23) than in Trechus beusti (Fig. 11, 21). The copulatory pieces are almost identical (Fig. 22, 24). Trechus beusti is larger, with elytra more oval and elytral striae less impressed (Fig. 4); Trechus pieltaini is smaller, narrower, and elongate with subparallel elytra and striae more impressed (Fig. 5).

Trechus brucki and Trechus bruckoides sp. n.do not have the peculiar hooked apex of the median lobe observed in the other species of the clade, but the apex is nevertheless strongly curved (Figs 25, 27).

The case of two further species remains doubtful: Trechus carrilloi was included by its descriptor in the uhagoni group especially because of the structure of the aedeagus, with an apex with an apical hook (Fig. 31). However, the secondary sclerotization of the ejaculatory duct is lacking in this species and it is characterized by a homogenous elytral pubescence which is present in other species of the area (Ortuño and Jimenez–Valverde 2011), but lacking in all the species of the Trechus brucki clade sensu novo. The presence of a hook at the apex of the aedeagus is also known in other Pyreneo–Cantabrian species like Trechus arribasi Jeanne, 1988, currently included in the Trechus fulvus group (Toribio 2001, Reboleira et al. 2010) or Trechus aubryi from Ariège. This character led its descriptor to include Trechus aubryi in the Trechus uhagoni group. Queinnec and Ollivier (2011) included the species in the Trechus angusticollis group. The species appears to be the sister species of Trechus distinctus (Fig. 34).

Finally, Trechus sharpi was included in the Trechus uhagoni group by Jeannel (1927), but the shape of the median lobe of the aedeagus and the copulatory piece, that shares some similarities with the Trechus bordei group, together with the lack of the sclerotization of spermiduct present in all the species of the Trechus brucki group sensu novo, cast doubts on its real phylogenetic affinities.

The species of the Trechus brucki clade are humicolous (grenieri, uhagoni), orophilous (grenieri, brucki, bruckoidessp. n.), or troglobitic/subterranean (bouilloni sp. n., pieltaini, beusti). We can notice a coincidence in the ecology of Trechini and Leptodirini with Basque–Pyrenean distribution: whereas the species are humicolous (or nivicolous for some Trechini) in the Pyrenees, the species occurring in the Basque country are mainly hypogean (Salgado et al. 2008, Ribera et al. 2010).

Biogeography of the Trechus brucki clade

If we use the standard mitochondrial mutation rate for insects of 2.3% divergence per Myr (0.0115 substitutions ⁄ site ⁄ Myr) (Brower 1994, Papadopoulou et al. 2010, Pons et al. 2010, Ribera et al. 2010), the isolation between the Trechus navaricus and Trechus brucki group seems to have occurred at the end of the Pliocene (Faille et al. 2011a). Pliocene climate was much warmer than the Present (Uriarte 2003): the interval between 3.3 Myr to 3 Myr was called Mid Pliocene warm Period, with an average temperature of about 3°C higher than at present and an annual average precipitation between 400 to 1000 mm higher than present. The transition to the Pleistocene (ca 2.7 Myr) is marked by the onset of marked climatic variability; the radiation of the Trechus brucki clade occured during the Pleistocene, following the rhythm of alternations of cold, warm/humid and dry periods that led to changes in biome composition (Salzmann et al. 2008).

Strong erosion leading to a deep excavation of Pyrenean valleys associated with climate variations led to the dispersal and diversification of the brucki clade. The main events are (Barrère 1963, Campos 1979, Serrat and Ventura 1993, Calvet 2004):

1. Persistence of the Ebro depression between the Basque–Pyrenean area and the Iberian central plateau. The persistence of the Ebro salty basin from the late Oligocene (25 Ma) until the late Miocene (6 Ma) isolated groups with an Iberian distribution from those with a Pyrenean or Basque–Pyrenean distribution. This flat and shallow lagoon area received the tributaries of the Ebro river, from Reinosa to the Mediterranean Sea.

2. Impact of Quaternary erosion on karst fragmentation. On the northern slope, the folds which have an east–west orientation are narrow and divided by north–south valleys. On the southern slope, orogenesis caused the formation of two folds with an east–west orientation (internal and external “sierras”) parallel to the axial chain. Similarly, Quaternary erosion separated these sierras by narrow north–south valleys. Near the Atlantic, these “sierras” meet with Basque folds which have a complex north–west/south–east orientation, divided by narrow north–south valleys, from Bilbao to Alsasua. Between Vitoria and Pamplona, these Basque “sierras” are separated by the Pre–pyrenean middle depression, a broad valley excavated by the Zadorra (westward) and Arakil (eastward) rivers (Fig 35). These rivers flow into the Ebro Basin, separating the northern massifs of Aralar, Urquilla and Gorbea from the southern Sierra of Urbasa–Andía. The hydrographic system was set mainly by significant erosion due to numerous glaciation cycles during the Pleistocene (2.5 Ma).

Our molecular study suggests that the brucki lineage could have originated in the area delimited by the northern sierras of Gorbea and Urquilla and the edge of the sierra de Andía. The sierras de Andía, Urbasa and Entzia form the exact border between the hypogean fauna of the Pyrenees and Iberia. North of this limit occur Trechus bouilloni sp. n., Troglorites breuili Jeannel, 1919 (Carabidae, Pterostichini) –Urbasa–Andía–Entzia, Aralar, Ernio and Pagoeta massifs, between the Deba and Urola rivers (Ortuño et al. 2010)–, Euryspeonomus eloseguii (Español, 1948), Bathysciola rugosa (Sharp, 1873), –Leiodidae, Cholevinae, Leptodirini which also belong to a clade of Basque–Pyrenean distribution (Ribera et al. 2010)– whereas south of this area (Sierras de la Demanda and Lóquiz, surrounding the Ebro basin) is characterized by a lack of Leptodirini and Troglorites. The only cave Coleoptera is Trechus schaufussi comasi (Basaula cave in Baríndano, south of Urbasa). Trechus schaufussi Putzeys, 1870 is a model of Iberian extensive distribution: it is widespread in the Iberian Peninsula, from Algarve in Portugal to Cantabria, Iberian Central System and the pre–pyrenean massif of Guara in Spain (Jeanne and Zaballos 1986, Zaballos and Jeanne 1994, Serrano 2003). This species is known to have separated early from Trechus sensu stricto (Faille et al. 2010a, 2011a).

Trechus bouilloni sp. n. has a subterranean lifestyle among the scree–covered northern slope (900 m) of Sierra de Andía, whereas the type locality of Trechus uhagoni is the Orobe doline (700 m), located at the eastern limit of Sierra de Urquilla. Early Pleistocene climate variations could have led to drastic changes in biome composition, limiting dispersal possibilities and leading to the isolation of the population of Trechus bouilloni sp. n. (potentially forestal), south of the Arakil River. One hypothesis could be that the hygrophilous species were colonizing high altitude or hypogean habitats during interglaciar warming as observed in other species of Coleoptera (Hernando et al. 1999, Faille et al. 2011b). These climate fluctuations might also have led to the western subterranean colonization of the two hypogean species, Trechus beusti (Sierra de Urquilla) and Trechus pieltaini (Sierra de Gorbea) while the group colonized the Pyrenean chain and diversified in numerous forms living in humid forests and alpine zones, from the Iraty Valley to the Ariège Basin. Migration toward East could have been possible along the small sierras of Tajonar and Labia, which link the Basque Mountains to the Pyrenees.

Trechus brucki lives in the alpine zone (above 1700 m) of the axial ridge from Pic d´Orhy to Col du Pourtalet, in the high Ossau Valley. On the north ridge, Trechus brucki can be encountered in the same biotopes, near snow tongues melting on scree-covered slopes, from Aspe to Gave de Pau Valleys. As for Trechus grenieri and Trechus uhagoni, both are mainly forestal and occur at lower altitude except for Trechus grenieri in the eastern part of the range (Mount Valier area). The Ariège Valley is the eastern limit of the group.

The distribution area of the Trechus brucki group coincides with the one of the Basque–Pyrenean Leptodirini clade (Fig 35). In the Pyrenees, both groups are made up of forestal, endogean, humicolous, lapidicolous or orophilous, but not hypogean, species. It is only in Basque relief, the western part of their distribution, that both groups include subterranean species. Regarding Leptodirini, the basal group of the Basque–Pyrenean clade is the Bathysciola schiodtei group (endogean/humicolous elements); its distribution area is extended from Ariège, Trechus mystica Fresneda & Fery, 2009 (France: Haute–Garonne and Ariège; Spain: Val d’Aran) to the Basque relief, Bathysciola breuili Bolívar, 1921 in Peña Gorbea or Bathysciola rugosa (Sharp, 1873) in Sierra de Urbasa and Urquilla. A high degree of troglobiomorphy is only found in some hypogean species of the Basque area: Aranzadiella Español, 1972 (basin of Deba River), Euryspeonomus Jeannel, 1919 (Aralar, Urbasa/Andía and Baztan Valley), Josettekia Bellés & Déliot, 1983 (Ernio and Aralar massifs), Nafarroa Fresneda & Dupré, 2010 (Kintoa Massif) and Speocharidius Jeannel, 1919 (between the Urola and Orio Rivers). In the Pyrenees, the species of the Trechus brucki clade are epigean, forestal (Trechus grenieri, Trechus uhagoni ruteri) or orophilous(Trechus brucki, Trechus bruckoides sp. n.). Pyrenean speciation events in the group are more recent and are probably closely related to late Pleistocene climatic changes, as already observed in alpine Trechus (Lohse et al. 2011). Troglobiomorphic features (depigmentation, microphthalmy) only occur in the hypogean Trechus beusti and Trechus pieltaini, both of them located in the Basque area. The two other species of this geographical area are located in wet and cold dolines (Trechus uhagoni) or subterranean environments (Trechus bouilloni sp. n.). Their general appearance (pigmented, well–developed eyes), similar to other epigean species, could be an indication of the recent colonization of this reduced habitat.

Figure 1.

Habitus of Trechus bouilloni sp. n. (Lizarraga pass).

Figures 2–3.

Habitus of 2 Trechus grenieri (grotte de l´Eglise) and 3 Trechus uhagoni (Orobe doline).

Figures 4–5.

Habitus of 4 Trechus beusti (Cueva de San Adrián) and 5 Trechus pieltaini (Cueva de Mairuelegorreta).

Figures 6–7.

Habitus of 6 Trechus bruckoides sp. n. (Montagnon) and 7 Trechus brucki (Jaout).

Figure 8–10.

Aedeagus in dorsal view of 8 Trechus bouilloni sp. n. (Lizarraga pass) 9 Trechus grenieri (Lapiaz de Lazur) and 10 Trechus uhagoni (Orobe doline). CP2, secondary copulatory piece.

Figures 11–14.

Aedeagus in dorsal view of 11 Trechus beusti (Cueva de San Adrián) 12 Trechus pieltaini (Cueva de Mairuelegorreta) 13 Trechus bruckoides sp. n. (Montagnon) and 14 Trechus brucki (Lac d’Anglas). CP2, secondary copulatory piece.

Figures 15–20.

Aedeagus in lateral view and detail of internal sac of 15, 16 Trechus bouilloni sp. n. (Lizarraga pass) 17, 18 Trechus grenieri (Eglise cave) and 19, 20 Trechus uhagoni (Orobe doline). CP2, secondary copulatory piece.

Figures 21–28.

Aedeagus in lateral view and detail of internal sac of 21, 22 Trechus beusti (Cueva de San Adrián), 23, 24. Trechus pieltaini (Cueva de Mairuelegorreta), 25, 26 Trechus bruckoides sp. n. (Montagnon) and 27, 28 Trechus brucki (Lac d’Anglas). CP2, secondary copulatory piece.

Figures 29–32.

29 Genital armature of the female of Trechus bouilloni sp. n. (Lizarraga pass) 30 Aedeagus in lateral view of Trechus sharpi (Cueva la Cuevona) 31, 32 Aedeagus in dorsal and lateral view of Trechus carrilloi (Bosque de Saja).

Figure 33.

Lectotype and paralectotype of Trechus brucki.

Figure 34.

Phylogram of Trechus of the brucki group obtained in RAxML, using the combined data matrix. Number in nodes: ML bootstrap (>50%) (see Material and Methods for details). In blue, the Trechus brucki group sensu novo. In purple, Trechus bordei group. In red: Trechus bouilloni sp. n.

Figure 35.

Distribution map of Trechus brucki group and related species. Material studied: symbols with cross.

Figure 36.

The MSS of Lizarraga pass (Navarra, Spain).

Figure 37.

Sinkhole area of the Plateau of Esturou (Hautes–Pyrénées, France).


We thank the collectors mentioned in Table 1 for providing material, M. Toribio and J.M. Salgado for the loan of material, I. Ribera and two referees for helping with their comments, T. Deuve and A. Taghavian (MNHN) for the loan of types and specimens of the T. brucki group and M. Balke (ZSM) who let us study the material of Trechini from the ZSM.

We are grateful to B. Jaeger (Museum für Naturkunde, Berlin) for providing information on the material from the Schaufuss collection and to J. Judge (University of California, Berkeley) for the linguistic revision of the manuscript. AF was supported by a postdoctoral Research Fellowship from the Alexander von Humboldt Foundation.

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Distribution of the Trechus brucki group sensu novo: Material studied and bibliographic records (Fig. 35, map). Material not studied indicated by (!)

Trechus brucki group sensu novo

Trechus beusti Schaufuss, 1863

Anophthalmus beusti Schaufuss 1863: 149

Spain: Guipúzcoa: 1. Zegama, Cueva de San Adrián (Schaufuss 1863; Bolívar y Pieltain and Jeannel 1921; Jeannel 1921, 1927; Jeanne and Zaballos 1986; Zaballos and Jeanne 1994; Serrano 2003; Ortuño and Marcos 2003); 4–VIII–2009, Fresneda leg., 1 ♀, voucher number label “ZSM_L199” (ZSM); 14–VIII–2009, Bourdeau and Fresneda leg., 1 ♂, voucher number label “ZSM_L200” (CAF); 31–XII–2009, Bourdeau leg., 4 exx. (CCB); 18–IV–2010, Bourdeau leg., 1 ex. (CCB); 2–V–2011, Bourdeau leg., 1 ♂ and 1 ♀ (CJF), 5 exx. (CCB). 2. Oñate, cave Iritegui (= Integui) (!) (Serrano 2003; Ortuño and Marcos 2003). 3. Oñate, cave Tortuga (!) (Serrano 2003; Ortuño and Marcos 2003).

Trechus bouilloni Faille, Bourdeau & Fresneda, sp. n.

See results

Trechus bruckoides Faille, Bourdeau & Fresneda, sp. n.

See results

Trechus brucki Fairmaire, 1862

Trechus brucki Fairmaire, 1862a: 548

Trechus politus Fairmaire, 1862b (nec politus Brullé, 1842): 578

Trechus planiusculus Fairmaire, 1862b (nec planiusculus Costa, 1858): 578

Trechus brucki microthorax Coiffait, 1952: 190

Trechus brucki pecoudi Colas & Gaudin, 1935: 248 n. syn.

Trechus brucki truilheti Coiffait, 1952: 189

Trechus brucki vandeli Coiffait, 1952: 189

France: Pyrénées–Atlantiques: 1. Val d’Ossau, les Eaux Bonnes (Fairmaire 1862a; Jeannel 1927, 1941). 2. Sommet du Pic d’Orhy, 1900 m (ssp. pecoudi, Colas and Gaudin 1935; Zaballos and Jeanne 1994; in Spain, Navarra, Macizo de Orhí after Serrano 2003); Pic d’Orhy, 5–1925, 1 ♂ and 1 ♀, coll. Nègre (MNHN); Basses Pyrénées, pentes du Pic d’Orry, 29–V–1936, G. Tempère / pierres, 1900–2000 m, 1 ex., coll. Nègre (MNHN), 1 ♂, coll. Coiffait (MNHN). 3. Pic de Jaout, dans la zone subalpine, vers 1500 m (!) (Jeannel 1941, Bonadona 1971). 4. Col de Mahourat, au fond de la vallée d´Ossau, près du col du Pourtalet, 1 ♀ (Coiffait 1952); Col du Pourtalet, Basses Pyr, pic de Maourat, alt 2000 m N°776, 28–9–1959, M. LAVIT (Holotype of Trechus brucki truilheti, coll. Coiffait, MNHN), 1 ♂ (Paratype of Trechus brucki truilheti, coll. Coiffait, MNHN). 5. Sesques, Lac Isabe, 10–8–1979, 8 exx., Bourdeau leg (CCB). Laruns, Pic de Sesques, bord de névé, 2300 m, 30–VI–2011, 1 ♀, voucher number label “ZSM–L446”, Bourdeau leg. (CAF). 6. Lescun, 1 ♂ (Coiffait 1952, Bonadona 1971). 7. Pic d´Anie, 1 ♀ (Coiffait 1952); Pic d´Anie, 1 ♀ bois de Braca, 15–VI–1951, Coll Coiffait (Holotype of Trechus brucki vandeli, coll. Coiffait, MNHN). 8. Pic d’Arlas, 1800 à 2000 m (!) (Jeanne 1984). 9. Col de la Pierre Saint Martin (!) (Jeanne 1984, in Spain after Zaballos and Jeanne 1994 and Serrano 2003). 10. Cirque d´Azuns, 1800 m (!) (Jeanne 1984). 11. Gourette, Lac d’Anglas, 20–8–2000, 6 exx., Bourdeau leg. (CCB). 12. Jaout, 15–8–1989, 1 ♂, Bourdeau leg. (CCB); 20–VI–1996, 1 ex., Bourdeau leg. (CCB); VIII–1981, 8 exx., Bourdeau leg. (CCB). 13. Col du Pourtalet, VI–2007, 3 exx., Bourdeau leg. (CCB). Pic d´Anéou, 2000 m, du col du Pourtalet au col d´Anéou, 2000 à 2100 m (Jeanne 1984). Pourtalet, Caperan d´Anéou, 5–VII–2011, 1 ♂, voucher number label “ZSM–L449”, 1 ♀, Bourdeau leg. (CAF). 7–1959, 2 exx, J. Aubry leg (coll. Coiffait, MNHN). 14. Pène Blanque, 2500 m (!) (Jeanne 1984). 15. Ossau, Pic de Gaziès, VII–2009, 1 ♀, voucher number label “ZSM–L190”, Bourdeau leg. (CAF). Hautes–Pyrénées: 16. Arrens, Pic du Gabizos, 2000 m, 10–VII–2010, 1 ♂, voucher number label “ZSM–L329”, 1 ♀, voucher number label “ZSM–L329bis”, Bourdeau leg. (CAF, ZSM), 1 ♀ (CAF). 17. Pic Granquet, ♂♂ et ♀♀ (Coiffait 1952); Pic Granquet, 1600 m, 5–9–1943, 8 exx. (Holotype and Paratypes of Trechus brucki microthorax, coll. Coiffait, MNHN); 1 ex, cotype, 1600 m, 5–9–1943, coll. Nègre (MNHN). Soum de Granquet, Lac d´Ourrec, en haute vallée de l´Esponne (!) (Queinnec and Ollivier 2011).

Spain: Huesca: 18. Candanchú (!) (Zaballos and Jeanne 1994). After Serrano (2003) in Spain, «Lérida» (mistake), Macizo de Aneu (!).

Trechus grenieri Pandellé, 1867

Trechus grenieri Pandellé, 1867: 147

Trechus bepmalei Jeannel, 1921: 176

Trechus despaxi Jeannel, 1922: 341

Trechus uhagoni prepyrenaeus Coiffait, 1974: 24

Trechus uhagoni aulaensis Aubry, 1981: 251

France: Hautes–Pyrénées: 1. Gazost, 1200 m, au bord des ruisseaux en forêt (Pandellé 1867; Jeannel 1927, 1941, Bonadona 1971), 1 ♀, coll. Fairmaire 1906 (MNHN). 2. Val de Lesponne, sous les amas de feuilles mortes en bordure immédiate des ruisseaux (!) (Bonadona 1971). 3. Barèges, 1 ♂, ex coll. Jeannel ex coll. Castelnau (MNHN). 4. Fréchet–Aure, résurgence de la Hèche, 14–V–2008, 1 ♂, voucher number label “ZSM–L13”, 1 ♀, Besson, Bourdeau & Faille leg. (CAF). 5. Nistos, Bas–Nistos, Grotte de l’Eglise (Bonadona 1971, Corbaz & Jauzion 1988); 2–III–1980, 2 exx., Bourdeau leg (CCB); 27–V–1945: 1 ♂, 17–VI–1945: 1 ♂, coll. Fourès (MNHN); 1 ♂, M. Bouillon rec. (CAF); VI–46, 4 exx., Bourgoin (MNHN); 15–XII–45, 3 exx., Bourgoin Colas (MNHN); 28 exx. (MNHN, coll. Coiffait). 6. Doline de la Bayelle de Gazave, 6 exx. (MNHN). Haute–Garonne: 7. Saint–Béat, Cap de Tus, 1200 m, près de la fontaine ferrugineuse au–dessus du col de Couret (Jeannel 1922, 1927, 1941, Bonadona 1971); été 1922, R. Despax, 3 exx., coll. R. Jeannel (MNHN). Saint Béat, Août 1922, 1 ex., coll. Despax in coll. Nègre (MNHN). 8. Boutx, forêt de Mourtis (Jeannel 1927, 1941, Bonadona 1971). Ft de Mourtis, St Béat, 1450 m, VIII–1926, 9 exx., coll. R. Jeannel, 1931 (MNHN). Ft de Mourtis, 5 exx., coll. Nègre (MNHN). 9. Arbas, 4–8–1980, 9 exx., Bourdeau leg. (CCB). 10. Val d’Espingo, 1800 m, au–dessus du lac d’Ôo (!) (Jeannel 1921, 1927, 1941). 11. Haute vallée du Lys, Superbagnères (Jeannel 1941, Bonadona 1971). Station de Superbagnères, 1650–1700 m, 3–X–1929, 1 ♂, Jeannel (MNHN). « Station de Superbagnères/ pierres 1700 m, 3 oct 1929 », 1 ♂, Gadeau de Kerville, coll. Nègre (MNHN). Ariège: 12. Rimont, Maison forestière, VII–1962 (Trechus uhagoni prepyrenaeus, Holotype ♂, coll. Coiffait MNHN). 13. Forêt d´Andronne, Le Bosc, vers 1000 m, 1 ex., 2 ♂♂ and 4 ♀♀ (Coiffait 1974). IV–1961, 2 ♂♂, Trechus uhagoni prepyrenaeus, paratypes; XI–1961, 3 ♀♀, paratypes (MNHN, coll. Coiffait). 14. Riverenert, vers 1100 m sur le versant Nord du col de la Crouzette, 2 ♀♀ (Coiffait 1974). La Crouzette, Sentenac de Serou, IX–1960, 2 ♀♀, coll. Coiffait (MNHN). 15. Lapiaz de Lazur, flanc NE du Mont Valier, VII–1978, 2 exx., Bourdeau leg. (CCB). 16. Port d’Aula, 2200 m (!) (Aubry 1981). Spain: Lérida: Port d’Aula (!) (Zaballos and Jeanne 1994; Serrano 2003).

Trechus pieltaini Jeannel, 1920

Trechus pieltaini Jeannel, 1920: 155

Spain: Álava: 1. massif of Gorbea (Serrano 2003), Cueva de Mairuelegorreta (Jeannel 1920, 1927, Bolívar y Pieltain and Jeannel 1921, Ortuño and Marcos 2003); 22–V–2011, 1 ♀, voucher number label “ZSM–L395”, Bourdeau leg. (CAF). 22–VII–2011, 4 ♂♂ and 4 ♀♀, Bourdeau leg. (CJF, CAF). 9.IV.1977, 2 exx., Garde leg. (coll. Lagar). 2. Cueva del Manantial (!) (Bolívar y Pieltain and Jeannel 1921). 3. Cueva de Arcegui (Monte Gorbea 1000 m, 30TWNI1963), T.M. Zuya–Zuia (!) (Ortuño and Marcos 2003). 4. Cueva de Sogusti–2 (Monte Gorbea 1000 m, 30TWNI1963), T.M. Zigoitia (!) (Ortuño and Marcos 2003). Vizcaya: 5. Cueva del Polvorino (= Polvorón) de Elorrea, Ceánuri en el macizo de Gorbea, a 1050 m sobre el nivel del mar (Español 1965, Zaballos and Jeanne 1994); 21–IX–1962, 1 ♀, Nolte leg., coll. Daffner (ZSM). 6. Ceánuri, Sima A–S–109 (!) (Zaballos and Jeanne 1994).

“Hisp”, “Alte Sammlung”, 3 ♂♂, 1 ♀ (ZSM). We found in the ZSM collection a specimen labelled “Asturien, collection Strasser” which is most probably an erroneous locality.

Trechus uhagoni Crotch, 1869

Trechus uhagoni Crotch, 1869: 14

Spain: Navarra: 1. Zegama, Alsasua (Crotch 1869) Cueva de Orobe (Jeannel 1927; Español 1965, Zaballos and Jeanne 1994; Serrano 2003); 1–VI–2004, 7 exx., Bourdeau leg. (CCB); 13–VII–2004, 1 ♂, voucher number label “MNHN–AF102”, Bourdeau leg. (MNHN), 1 ♀ (CAF); 2–V–2009, 3 exx. (CJF, CAF); 1 ♂, voucher number label “ZSM–L161” Bourdeau and Fresneda leg. (CAF).

We found 4 specimens labelled «Andara Escalera» (MNHN). A locality called Endara exists in SE of Oiartzun (Guipúzcoa). We cannot exclude that the species also occurs in this area. It is also quoted in Guipúzcoa, Macizo de Izarraitz, Ekain (Galán 2003) but the specific attribution of these Trechus should be verified.

Trechus uhagoni ruteri Colas, 1935

Trechus uhagoni ruteri Colas, 1935: 253

France: Pyrénées–Atlantiques: 1. Larrau, Cañon d´Holçarté, pont d´Amuby, 7–1934, G. Colas (Holotype, aedeagus not with the specimen (MNHN) (Colas and Gaudin 1935, Jeannel 1941, Bonadona 1971). 2. Larrau, Bois de Saint Joseph (!) (Jeanne 1984). 3. Forêt d´Iraty (Jeanne 1984); Iraty, 21–IX–1949, 6 exx., H. Coiffait (MNHN, coll. Coiffait). 4. Col de Bentarté, près du Mont Urculo, 5–1925, 4 ♂♂ and 1 ♀♀, coll. Jeannel (MNHN) (Jeannel 1941, Bonadona 1971, Jeanne 1984).

Spain: Navarra: 5. Aoiz, Acueducto de Orbaiceta (Español 1977). 6. Orbaiceta, Bosque de Irati (Jeanne and Zaballos 1986, Zaballos and Jeanne 1994). 3. Espinal, puerto de Ibañeta (Zaballos and Jeanne 1994). 7. Col de Roncevaux, juillet 1934, 2 ♀♀, L. & A. Gaudin (MNHN). 5–1925, 2 exx. (Coll Nègre in MNHN). 8. Entrada C. de Espinal, C. Bolívar, 2 ♂♂ (MNHN). 6. Peña Escaori, 1600 m, 5–1925, 1 ♂, Gaudin (MNHN).

Species of uncertain phylogenetic affinities

Trechus carrilloi Toribio & Rodríguez, 1997

Trechus (Trechus) carrilloi Toribio & Rodríguez, 1997: 283

Spain: Cantabria: 1. Campoo de Cabuérniga, Bosque de Saja, UTM 30TUN967728 (Toribio and Rodríguez 1997; Serrano 2003); type series (!) (Toribio and Rodríguez 1997): Holotype ♂, 23–VIII–1997, M. Toribio leg. (CMT); paratypes (CMT, MNCN, col. Carabajal, col. García and col. Rodríguez): 4–X–1997, 11 ♂♂ and 9 ♀♀, J. García leg.; 4–X–1997, 3 ♂♂ and 4 ♀♀, F. Rodríguez leg.; 31–X–1997, 2 ♂♂, M. Toribio leg.; 31–X–1997, 2 ♂♂ and 2 ♀♀, F. Rodríguez leg.; 21–XI–1998, 2 ♂♂, F. Rodríguez leg. Same locality, 27–VIII–2001, 1 ♂, Toribio leg. (CJF).

Trechus sharpi Jeannel, 1921

Trechus sharpi Jeannel, 1921: 179

Spain: Cantabria: 1. Santander, Población, sur le mont Hijedo, au sud–est de Reinosa (Jeannel 1921, 1927; Zaballos and Jeanne 1994). 2. Santander, Collection Strasser, 1 ♂ (ZSM). 3. Puerto de San Glorio (Zaballos and Jeanne 1994). 4. Pico Tres Mares (Zaballos and Jeanne 1994). 5. Montes cantábricos orientales (Serrano 2003). 6. Sejo (T.M.Valdaliga), Cva. La Mina, 20–XII–2003, 2 ♂♂, C.G. Luque leg. (Col. Salgado). 7. Quijas, Cueva la Cuevona, 13–IX–1995, 1 ♂, C.G. Luque leg. (Col. Salgado). 8.Trechus cantaber”, Monts Cantabriques, Sierra de la Sagra, 3 exx., coll. de la Cruz in coll. Coiffait (MNHN).