Ann. soc. entomol. Fr. (n.s.), 2011, 47 (1–2) : 21-32 ARTICLE

Taxonomic notes on Trechini and description of a new hypogean species from the Iberian Peninsula (Coleoptera: Carabidae: ) Vicente M. Ortuño (1) & Alberto Jiménez-Valverde (2) (1) Departamento de Zoología y Antropología Física. Facultad de Biología, Universidad de Alcalá, E-28871 - Alcalá de Henares. Madrid, Spain (2) Natural History Museum and Biodiversity Research Center, Th e University of Kansas, Lawrence, Kansas 66045, USA

Abstract. In this contribution, a new pubescent species for the Iberian Peninsula is described, and the use of some morphological traits in the systematic of Trechini are discussed. Micropubescence is common in endogean and troglobite species which are closely related with the “terrestrial-phreatic habitat”. It probably enhances survival in swampy environments through the retention of air bubbles in immersion conditions and promoting fl otation. Neither this trait, anophthalmy nor inner sac of aedeagus without sclerotized pieces can be used to support supraspecifi c taxa. Thus, we propose the formal synonymy of Hydrotrechus with Trechus [Hydrotrechus syn. nov. of Trechus]. To avoid secondary homonymy, we propose the name Trechus triamicorum nom. nov. for Hydrotrechus cantabricus Carabajal, García & Rodríguez 1999 [already occupied by Trechus cantabricus Jeannel 1927, synonym of Trechus distigma Kiesenwetter 1851]. We also propose the synonymization of the subgenus Trichapoduvalius with Apoduvalius [Trichapoduvalius syn. nov. of Apoduvalius]. The new hypogean species with pubescent elytra, Trechus cifrianae sp. nov., is described. The morphology of the species suggests that it probably inhabits the Mesovoid Shallow Substratum (MSS) or the fi ssure network (mesocaverns and microcaverns) of the cave where the only individual was found. The structure of internal female genitalia suggests a close relationship between T. cifrianae sp. nov. and T. carrilloi Toribio & Rodríguez 1997. Résumé. Notes taxonomiques sur les Trechini et description d’une nouvelle espèce hypogée de la Péninsule Ibérique (Coleoptera : Carabidae : Trechinae). Dans cette contribution, une nouvelle espèce pubescente de Trechus est décrite de la Péninsule Ibérique, tandis que l’utilisation de certains caractères morphologiques de la tribu des Trechini est discutée. La micropubescence est fréquente chez les espèces endogées et troglobies ce qui est à mettre en relation avec l’habitat “terrestre- phréatique”. Ce caractère augmente probablement la survie dans les envirronnements marécageux du fait de la rétention des bulles d’air en immersion qui améliorent la fl ottaison. Ni ce trait, ni l’anophtalmie, ni le sac interne des édéages sans pièce sclérifi ée ne peuvent être utilisés pour caractériser les taxons supraspécifi ques. Dès lors, nous proposons de mettre en synonymie Hydrotrechus avec Trechus [Hydrotrechus syn. nov. of Trechus]. Pour éviter les homonymies secondaires, nous proposons le nom Trechus triamicorum nom. nov. pour Hydrotrechus cantabricus Carabajal, García & Rodríguez 1999 [préoccupé par Trechus cantabricus Jeannel 1927, synonyme de Trechus distigma Kiesenwetter 1851]. Nous proposons aussi la mise en synonymie du sous-genre Trichapoduvalius avec Apoduvalius [Trichapoduvalius syn. nov. de Apoduvalius]. Une nouvelle espèce hypogée à élytres pubescents est décrite : Trechus cifrianae sp. nov. La morphologie de cette espèce suggère qu’elle habite probablement le “Mesovoid Shallow Substratum” (MSS) ou les réseaux de fi ssures (mésocavernes et microcavernes) de la cave d’où le seul spécimen connu provient. La structure des génitalia internes femelles suggère une relation étroite entre T. cifrianae sp. nov. et T. carrilloi Toribio & Rodríguez 1997. Keywords: Taxonomy, Systematic, Coleoptera, Carabidae, Iberian Peninsula.

arabidae are extremely diversifi ed coleoptera in- dwell these aphotic and moist habitats, playing an im- Chabiting almost all terrestrial ecosystems (Lövei portant role in the equilibrium of subterranean eco- & Sunderland 1996). Subterranean endogean cham- systems. Th e subfamilies Pterostichinae and, mainly, bers, hypogean environments (Mesovoid Shallow Sub- Trechinae, contain the lineages with the greatest di- stratum – MSS – and bedrock fi ssure network) and the versifi cation in hypogean habitats, exhibiting notice- transitional zone are among the cryptic habitats where able morphological and physiological adaptations (see Casale et al. 1998). Trechinae contains lineages that this family is well represented (Ortuño 2002). have suff ered an intense troglobiomorphy evolution Carabids showing predatory and scavenger behaviours (Dethier & Hubart 2005) and, so, they have a body shape that lies far from the epigean archetype. Th is results in the so called “aphaenopsian” morphology, E-mail: [email protected], [email protected] i.e., depigmented, anophthalmic and graceful mor- Accepté le 31 mai 2010 photypes, with a tendency to show false physiogastry

21 V. M. Ortuño & A. Jiménez-Valverde

(pseudophysiogastry: Dethier & Hubart 2005), long some of them closely related to Aphaenops as, for ex- appendages and a high number of sensory receptors. ample, Hydraphaenops Jeannel 1926. It is also present Th is morphology takes the name from the genus in Asiatic hypogean species from several genera and, Aphaenops Bonvouloir 1861, where the most genu- less frequently, in species from America and Oceania ine representatives are included. Th is corporal shape (see Casale et al. 1998). Until now, Trechini stylization is exhibited by many hypogean Trechini in Europe, shows the maximum exponent in the hypogean species

Table 1. Examined material.

Phyletic series Genus - species (sensu Casale et al. 1998) Trechus Clairville 1806 (57 studied species): T. fulvus Dejean 1831; T. lallemantii Fairmaire 1859; T. alicantinus Español 1971; T. martinezi Jeannel 1927; T. beltrani Toribio 1990; T. torressalai Ortuño & Arillo 2005; T. breuili Jeannel 1913; T. pisuenensis Ortuño & Toribio 2005; T. barratxinai Español 1971; T. diecki Putzeys 1870; T. escalerai Abeille 1903; T. gloriensis Jeanne 1971; T. apoduvalipenis Salgado & Ortuño 1998; T. saxicola Putzeys 1870; T. beusti (Schaufuss 1863); T. pieltaini Jeannel 1920; T. pecignai Toribio 1992; T. arribasi Jeanne 1988; T. carrilloi Toribio & Rodríguez 1997; T. distinctus Fairmaire & Laboulbène 1854; T. pyrenaeus Dejean 1831; T. latebricola Kiesenwetter 1850; T. sharpi Jeannel 1921; T. bordei Peyerimhoff 1909; T. jeannei Sciaky 1998; T. planipennis Rosenhauer 1856; T. kricheldorffi Wagner 1913; T. grenieri Pandellé 1867; T. suturalis Putzeys 1870; T. distigma Kiesenwetter 1851; T. ceballosi Mateu 1953; T. barnevillei Pandellé 1867; T. tingitanus Putzeys 1870; T. cuniculorum Méquignon 1921; T. schaufussi Putzeys 1870; T. “Trechus” quadristriatus (Schrank 1781); T. obtusus Erichson 1837; T. fl avocircumdatus Jeannel 1922; T. fl avocinctus Jeannel 1922; T. fl avolimbatus Wollaston 1863; T. minioculatus Machado 1987; T. umbricola Wollaston 1854; T. picoensis Machado 1988; T. terceiranus Machado 1988; T. longobardus Putzeys 1870; T. brembanus Focarile 1949; T. insubricus K. Daniel & J. Daniel 1898; T. magistrettii Focarile 1949; T. priapus K. Daniel 1902; T. amplicollis Fairmaire 1859; T. putzeysi Pandellé 1867; T. varendorffi Sainte-Claire Deville 1903; T. gracilitarsis K. Daniel & J. Daniel 1898; T. cardioderus Putzeys 1870; T. longulus K. Daniel & J. Daniel 1898; T. marginalis Schaum 1862; T. (Epaphius) secalis (Paykull 1790) Anchotrechus Jeannel 1927 (monobasic genus): A. punctipennis Jeannel 1927 Hydrotrechus Carabajal, García & Rodríguez 1999 (monobasic genus): H. cantabricus Carabajal, García & Rodríguez 1999 Geotrechus Jeannel 1919 (1 studied species): G. orpheus (Dieck 1869) Aphaenops Bonvouloir 1861 (5 studied species): A. jeanneli (Abeille 1905); A. ochsi L. Gaudin 1925; A. leschenaulti Bonvouloir 1862; A. orionis Fagniez 1913; A. (Cerbaphaenops) cerberus (Dieck 1869) Hydraphaenops Jeannel 1926 (1 studied species): H. vasconicus (Jeannel 1913) Paraphaenops Jeannel 1916 (monobasic genus): P. breuilianus (Jeannel 1916) “Aphaenops” Sardaphaenops Cerruti & Henrot 1956 (1 studied species): S. supramontanus Cerruti & Henrot 1956 Doderotrechus Vigna Taglianti 1968 (1 studied species): D. ghilianii (Fairmaire 1859) Boldoriella Jeannel 1928 (1 studied species): B. carminatii (Dodero 1917) Apoduvalius Jeannel 1953 (5 studied species): A. leonensis Salgado & Ortuño 1998; A. alberichae Español 1971; A. naloni Salgado 1993; A. champagnati Salgado 1991; A. anseriformis Salgado & Peláez 2004 “Iberotrechus” Iberotrechus Jeannel 1920 (monobasic genus): I. bolivari (Jeannel 1913) “Trechoblemus” Trechoblemus Ganglbauer 1891 (1 studied species): T. micros (Herbst 1784) Duvalius Delarouzée 1859 “Duvalius” (7 studied species): D. casellii (Gestro 1898); D. brucki (Piccioli 1870); D. carantii (Sella 1874); D. taygetanus Casale 1979; D. comottii Casale, Giachino, Vailati & Vigna Taglianti 1966; D. berthae (Jeannel 1910); D. lencinai Mateu & Ortuño 2006 “Pheggomisetes” Pheggomisetes Knirsch 1923 (1 studied species): P. buresi (Knirsch 1923) Neotrechus J. Müller 1913 (1 studied species): N. suturalis (Schaufuss 1864) “Neotrechus” Orotrechus J. Müller 1913 (1 studied species): O. muellerianus (Schatzmayr 1907); O. vicentinus (Gestro 1907) “Typhlotrechus” Typhlotrechus J. Müller 1913 (1 studied species): T. bilimeki (Sturm 1847)

22 Taxonomic notes on Trechini and a new species

Giraff aphaenops clarkei Deuve 2002, from southern Results China (Deuve 2002). In the Iberian Peninsula we can fi nd aphaenopsian morphologies not only in the genus Aphaenops and Hy- Trechus cifrianae sp. nov. draphaenops, but also in Paraphaenops Jeannel 1916 and, (Figs. 1–9, 11 and 16c) to a lesser extent, in Geotrechus Jeannel 1919, Apoduval- Material examined. Holotype: 1 ♀, “Cueva del Pis” cave, 310 ius Jeannel 1953 and some species of Duvalius Dela- m elevation, UTM-MGRS=30TVN29 (Penilla, Santiurde rouzée 1859. Nevertheless, not all subterranean Trechi- de Toranzo, Cantabria - Spain), 19.VIII.2004, V.M. Ortuño nae species show aphaenopsian shapes, and species with more robust shapes, relatively shorter appendages and ocular traces can be found; Iberian examples are Duval- ius lencinai Mateu & Ortuño 2006, Hydrotrechus can- tabricus Carabajal, García & Rodríguez 2000 and some species of Trechus Clairville 1806. Th e Trechus phyletic line (sensu Casale & Laneyrie 1982) is probably one of the most interesting due to its high specifi c diversity, including more than 600 Trechus species (Moravec et al. 2003) (a mainly holartic genus, Casale et al. 1998). Although most of them are epigean (edaphic life-style), hypogean species can also be found in this genus. Th is fact implies that several degrees of subterranean adapta- tions can be found in the same taxa, and it is illustrated in the Iberian fauna: from the 51 known species of Tre- chus (Serrano 2003; Ortuño & Toribio 2005; Ortuño & Arillo 2005), 19 show subterranean habits (endoge- an and hypogean), mainly inhabiting caves and abysses (Ortuño & Arillo 2005: 3484). Th e discovery of a new pubescent Trechus species in the “Cueva del Pis” cave (Cantabria, Spain), with mor- phological characteristics typical of subterranean and maybe amphibian habits, indicates, without doubt, a hypogean life. Together with the description of this new species we review some morphological traits of Trechini and their use in the systematic of the taxa. Material and methods Th e female genitalia was studied using the following procedure. Th e terminal abdominal segments of the female were dissected and placed in a saturated solution of KOH for 12 h. Th en, they were washed in Scheerpeltz’s solution (see Ortuño et al. 1992: 148) and opened dorsally to check the alkaline digestion. Staining was carried out with Chlorazol black E® in aqueous solution for 30 seconds and the excess dye was removed by washing in KOH and the structures washed again in Scheerpeltz’s solution. Th e dissected structures were placed in a watchglass with Scheerpeltz’s solution under a stereomicroscope with a drawing tube. Finally, the female genital preparations were included in DMHF and put on an acetate sheet. In the keys to species, an index is used to inform about the relative length of antennae [LRA = LA/LC], where LA is antennae length and LC is body length (from tip of mandible to elytron-apex). To complete the taxonomic study of the new species of Trechus we have studied the expression of some of its more relevant Figure 1 characteristics in other species of the genus. Additionally, spe- Trechus cifrianae sp. nov.: holotype. Right mesothoracic leg absent. Scale: cies from other genus present in Western Europe have also been 1 mm. studied (see tab. 1).

23 V. M. Ortuño & A. Jiménez-Valverde leg., deposited in the Department of Zoology and Physical with pronotum. Mandibles, labial and maxillary pieces as Anthropology – V.M. Ortuño (DZAF-UA/VMO), Universidad typical in the genus. Labial tooth grooved, truncated. Deep de Alcalá, Alcalá de Henares (Madrid, Spain). frontal sulci go round ocular area and extend towards clypeus. Etymology. Th is new species is dedicated to our friend Dra. Temples pubescent, strongly convex and long due to ocular Blanca Cifrián, professor and researcher in the Department regression. Non-functional eyes, reduced to small scars located in the front of ocular area. Antennae long (1/3 of elytra length), of Zoology and Physical Anthropology of the Universidad de st th Alcalá (Spain). pilose from 1 to 11 antennomeres. Most of the antennomeres are 2.5-3 times as long as wide. Cephalic chaetotaxy: one setae Diagnosis (Fig. 1). Length: 4.5 mm (from tip of mandible in the sulcus of each mandible; six setae on labrum; four setae to elytron-apex). Without developed wings. Anophthalmic on clypeus; two pair of supraocular setae (bilateral disposition), (small ocular scars). Depigmented (testaceous) and partially the anterior pairs slightly behind the ocular scar. Pronotum micropubescent (well visible in elytra and temples) integument. (Fig. 1): Convex, wider than long (length/width = 0.7), Pronotum trapezoid, wider than long, sides curved and slightly maximum width just in front of the middle line. Basal margin sinuous just before hind angles; these are sharp and almost right. rectilinear and slightly wider than anterior margin. Lateral Elytra sub-parallel with round humeral margin; deep and well- margin regularly curved and slightly sinuous just before hind defi ned striae nearly to apex. Head (Fig. 1). Almost as wide as angles; these are sharp and almost right. Lateral sulcus deep long (without considering mandibles); robust in comparison and narrow. Central sulcus pronounced. Basal foveae wide, deep and without micropuncture. Pronotal chaetotaxy: two

Figures 2–6 Figures 7–9 Trechus cifrianae sp. nov. 2, VI and VII sternites; 3, VIII ventrite; 4, VIII Trechus cifrianae sp. nov. 7, female external genitalia in ventral view; terguite; 5, detail of the setation of the VIII terguite; 6, IX terguite. Scale: 8, spermatheca in ventral view; 9, spermatheca in dorsal view. Scale: 7: 2: 0.5 mm; 3, 4 and 6: 0.3 mm; 5: 0.05 mm. 0.1 mm; 8 y 6: 0.2 mm.

24 Taxonomic notes on Trechini and a new species setae (bilateral disposition), one anterior seta in the fi rst quarter Anchotrechus (see micropuncture of A. punctipennis - and one posterior seta next to hind angle. Elytra (Figs. 1 and Fig. 16a-). Recently, a microphthalmic Trechus from 16c). Wide, glaringly sub-parallel, maximum width in the middle. Rounded-shoulders. Basal margin short and oblique. the MSS was discovered in the north of the Iberian Well-defi ned striae clearly stippled in their whole length. 8th Peninsula, Trechus carrilloi Toribio & Rodríguez 1997. stria well-separated from lateral sulcus. Apical striola deep and Th e most unique trait of this species, which is lacking linked to the end of the 5th stria. Interstriae slightly convex with puncture pilose in line. Lateral sulcus wide. Elytral chaetotaxy: scutellar pore at the beginning of 1st strias; discal setae in 3rd stria (anterior seta slightly behind the basal sixth and posterior seta in half ); umbilical series typical of the genus (humeral area with four equidistant setae and apical area with four setae, two anterior and two posterior). Legs (Fig. 1). As typical in the genus. Dorsal external margin of foretibia with a fi ne longitudinal sulcus. Aedeagus. Unknown. Female abdomen (Figs. 2–6). Harpalidian model (sensu Deuve 1988). VII sternite with four setae next to posterior margin (Fig. 2). VIII tergite (VIII mediotergite sensu Deuve 1993) with anterior concavity surrounded by laminated apophysis (Figs. 4–5). IX tergite (IX mediotergite sensu Deuve 1993) narrow and poorly sclerotized (Fig. 6). VIII ventrite (Fig. 3) wider than long, slightly sclerotized and hidden below VII sternite. Female external genitalia (=genital shield) (Fig. 7). Formed by dimerous IX gonopods (gonocoxite and gonosubcoxite), and IX laterotergites. Gonocoxite unguiform with 2 thorn- shaped setae on its dorsal surface, one near the internal edge and the other in the middle of the gonocoxite; small groove near apex with a fi ne sensorial seta. Gonosubcoxite as long as wide, globe-shaped, with three thorn-shaped setae (basal seta larger) in internal margin and another three setae (fi ner than the other three) in basal margin. Wing-shaped, slightly sclerotized (except external and anterior margins) IX laterotergite with one group of setae over basal margin (approximately 10) and one more internal group (approximately 6). Female internal genitalia (=spermathecal complex) (Figs. 8–9). Membranous; tubular-shaped vagina-bursa ending in spermatheca sacciform with densely folded walls (slightly auricle-shaped). Short blind prolongation in distal end of bursa. Long odd oviduct, narrow in the basal part (in contact with bursa invagination), dilated and with microfringes in distal half.

Discussion

Pubescence in the genus Trechus Although Jeannel (1927) pointed out that Trechus species show glabrous integument, this is not a general characteristic of the genus, as revealed by Israelson & Palm (1979). Th ese authors observed pubescence (rows of tiny setae) on the elytra of Trechus fortunatus Jeannel 1927 and Trechus uyttenboogaarti Jeannel 1936. Since then, and with the increase in the taxonomic knowledge of the genus, this characteristic has been confi rmed in other Macaronesian species (Machado 1992): Trechus machadoensis Franz 1984, Trechus laureticola Jeannel 1936 and Trechus gomerensis Figures 10–12 Franz 1986. Another species, Trechus cabrerai (Jeannel Antennae of three species of pubescent Trechus: 10, T. triamicorum nom. 1936), shows a very conspicuous pubescence covering nov. (= Hydrotrechus cantabricus); 11, T. cifrianae sp. nov.; 12, T. carrilloi. the elytra as well as the pronotum; this trait led Jeannel Scale: 0.5 mm. (1936) to incorrectly ascribe this species to the genus

25 V. M. Ortuño & A. Jiménez-Valverde in the rest of the Iberian species, is a row of microsetae of T. fortunatus is imperceptible (microsetae 10 μ long). in every interstria that gives a pubescent appearance to However, other species as, for example, T. carrilloi or T. the elytra (Toribio & Rodríguez 1997). cifrianae sp. nov., show elytral interstrias with distinct It is well-known the existence of pubescent as well micropuncture (Figs. 16b,c), a trait that is intimately as glabrous integuments in species pertaining to the related to the presence of a well-developed pubescence same genus in Trechini, as in Aphaenops, Geotrechus, (easily visible with grazing light). Th e expression of this Duvalius, Antoinella Jeannel 1937, among others, and character has been considered and ancestral trait in among the Iberian endemisms, as in Apoduvalius. Th us, Trechini (Jeannel 1936; Carabajal et al. 1999, among we have to be cautious when interpreting this morpho- others). However, in most Trechus species this trait is logical trait as, even though it can help to diff erentiate in recession, being reduced the size of the micropunc- certain species, it can hardly characterize a lineage with ture and, consequently, of pubescence, until its disap- generic status. As pointed out by Machado (1992), we pearance. Th us, glabrous integuments can retain mi- have also to change our concept of the genus Trechus cropuncture in most cases, even almost imperceptible and recognize a small number of more or less pubes- (Figs. 16e-g). cent species, including the new species that is described in the present article, Trechus cifrianae sp. nov. We also question the taxonomic identity of Hydrotrechus Cara- bajal, García & Rodríguez 2000 and Trichapoduvalius Vives 1976. We recommend transferring Hydrotrechus cantabricus, monotypic species, into Trechus, adding in this way one more pubescent species to the genus. A detailed study of several Trechus species reveals that pubescence is not just a qualitative character but also a quantitative one, both in the extension of the pubescence and in the length of the microsetae. For example, in words of Machado (1992), the pubescence

Figures 13–14 Figure 15 Aedeagus and detail of inner sac of two Trechus species: 13, T. triamicorum Detail of the pronotum and base of the elytra of Trechus triamicorum nom. nom. nov. (= Hydrotrechus cantabricus); 14, T. carrilloi. Scale: 0.1 mm. nov. (= Hydrotrechus cantabricus). Scale: 0.5 mm.

26 Taxonomic notes on Trechini and a new species

It seems evident that in Trechus (and in other and, in another, to describe a new genus and species Trechini) micropuncture and micropubescence are (Hydrotrechus) of Trechini. Accordingly to Carabajal et correlated traits (= linked by ontogeny) and, from a al. (1999), “Hydrotrechus se separa del género Trechus, cladistic point of view, they should be considered por varios e importantes caracteres: ausencia total de the same character. Otherwise, they would provide ojos, punteado y pubescencia elitral y edeago sin pieza redundant information aff ecting the independence copulatriz” (sic). Next, we will explain that the presence principle and altering results (Margaría & Lanteri of these traits, neither isolated nor combined, does not 2006). justify the proposal and maintenance of the genus Micropubescence is a common trait in endogeous Hydrotrechus. and cave-dwelling Trechini species (Laneyrie 1974), Ocular regression and, in the last term, anoph- closely related with the “terrestrial-phreatic habitat” thalmy, is a phenomenon that, together with other (sensu Jeannel 1926: 80). Rock fi ssures soaking in traits, helps to discriminate species in Trechini. How- water are the preferred habitat of terrestrial cave- ever, as it is a convergent trait – present in species with- dwellers (Jeannel 1926: 82) such as Hydraphaenops out a shared ancestor –, it must be considered with or of hypogean species such as the MSS inhabitant caution to avoid “taxonomic noise” in phylogenetic re- Hydrotrechus (Carabajal et al. 1999). Also, other constructions. Th us, the anophthalmy of H. cantabri- hygrophile Trechini inhabiting swampy terrains cus (it only shows a small ocular scar) cannot justify maintain the pubescence, such as Perileptina species the existence of the genus Hydrotrechus. Furthermore, sensu Moravec et al. (2003) (or Perileptini sensu Casale though it is more infrequent in Trechus than in Du- & Laneyrie 1982) from the genera Neoblemus Jeannel valius, several Iberian Trechus species show a notable 1923; Apoplotrechus Alluaud 1915 and Perileptus ocular regression, (Abeille 1903; Toribio 1990; Toribio Schaum 1860. Pubescent species can also be found in & Rodríguez 1997; Salgado & Ortuño 1998; Ortuño the Anillina (Bembidiini), inhabitants of the swampy & Toribio 2005), some of them showing a small area endogeous environment. Pubescence can retain of depigmented ommatidia (Trechus escalerai Abeille air bubbles in immersion conditions and promote fl otation in sudden fl ooding situations (Carabajal et al. 1999), thus explaining the amphibian life style and the capacity to inhabit subterranean (hypogean/endogean) environments of these carabids. The case of Hydrotrechus cantabricus At the end of the 20th century, in the Fagus sylvatica forest of Saja (Cantabria, northern Iberian Peninsula), an edaphic talus with well-developed A and B horizons and a C1 horizon (Mesovoid Shallow Substratum – MSS) of slate schist, was prospected. Th rough several survey campaigns, diverse Trechini species were found (among other ground species) living in syntopy. However, it has been confi rmed that these species inhabit diff erent depth levels and, thus, they probably occupy diff erent edaphic niches. Trechus distigma Kiesenwetter 1851 and the novelty Trechus jeannei Sciaky 1998 are common inhabitants of the A horizon. Another two species were also discovered in that survey inhabit the deepest horizons: Trechus carrilloi inhabits B and C1 horizons, and Hydrotrechus cantabricus inhabits exclusively C1 horizon. Both subterranean species show odd traits: pubescence (very Figure 16 conspicuous in the elytral interstrias and accompanied Detail of the and striae/interstriae and puncture of the elytra is some Trechini species: a, Anchotrechus punctipennis; b, Trechus carrilloi; c, Trechus cifrianae of micropuncture) (Figs. 15 and 16b,d) and ocular sp. nov.; d, Trechus triamicorum nom. nov. (= Hydrotrechus cantabricus); regression. However, these characters, together with e, Trechus fulvus; f, Trechus fl avocinctus; g, Trechus quadristriatus. Scale: 0.4 other ones, have been interpreted in diff erent ways, mm. being in one case used to describe a new Trechus species

27 V. M. Ortuño & A. Jiménez-Valverde

1903; Trechus beltrani Toribio 1990; Trechus carrilloi its male genitalia was used as a character to establish it. Toribio & Rodríguez 1997; Trechus apoduvalipenis Sal- It is worth to mention that there exists other species of gado & Ortuño 1998 and Trechus pisuenensis Ortuño the genus Duvalius s.str. with the pieces of the inner & Toribio 2005) while others showing ocular scars sac slightly sclerotized as, for example, the three species with nearly no sign of ommatidia (Toribio 1992) (Tre- of the Duvalius fodori-group (sensu Guéorguiev 2007): chus pecignai Toribio 1992 and the recent discovery, D. fodori Scheibel 1937; D. strupii Scheibel 1937 and Trechus cifrianae sp. nov.). D. vignai Casale 1983. Despite being included in a dif- Regarding the micropuncture of the integument, ferent subgenus, Guéorguiev (2007) call attention to intimately related to pubescence (mainly elytral their close relation with Platyduvalius. Although most but not exclusive), we have seen that, in Trechini, of the species belonging to the genus Trechus have scle- it is present in many clades (including Trechus). rotized anisotopic copulatory pieces, a small number of Presumably, it is an ancestral character that has been species have aedeagus without such structures. In the preserved (reverted in some cases) until now making latter case, the inner sac can have scales (for example, in possible the “amphibious-hypogean” life style. Th us, it Trechus alpicola and in species belonging to the Trechus is a plesiomorphic trait and it should not be used to lederi-group) (Jeannel 1927; Belousov & Kabak 1993) support the genus Hydrotrechus. or can be hyaline (for example, in Trechus brevicorpus Finally, the taxonomic value of the absence of scle- Belousov & Kabak 1993) (Belousov & Kabak 1993). rotized copulatory pieces in the inner sac (Fig. 13) Th ese are just an example of the diversity of structural has to be discussed. Although this trait is a good tax- expressions of the inner sac of the aedeagus in the tribu onomic tool, it must be evaluated if it is useful and Trechini. Th us, it is stated that the absence or even the suffi cient to defi ne a genus in Trechini. Th e biological presence of slightly sclerotized structures in the inner signifi cance of these pieces has been questioned; Avon sac of the aedeagus does not represent other thing that (1997: 31) pointed out that they may not be vital for a useful trait to diff erentiate species from a taxonomic reproductive activity. Th ere are several taxa in Trechini point of view. As follows, if according to all characters that lack sclerotized pieces, with a membrane or hya- a species can be placed into an existing genus, in no line sac (as in Sardaphaenops Cerruti & Henrot 1956) way the absence of sclerotized pieces in the inner sac or even covered by spines (for example, taxa from the can be used to propose a new genus. Th is is the case of Duvaliomimus phyletic series, sensu Casale & Laneyrie Hydrotrechus cantabricus, whose morphological charac- 1982) or scales (for example, taxa from the Lasiotrechus teristics place it inside the genus Trechus; the singular- phyletic series, sensu Casale & Laneyrie 1982). Anyway, ity of the inner sac of the aedeagus must be interpreted these singularities of the aedeagus are a complement to as a synapomorphy, a plausible and well-known trait in other morphological traits (setation included) which this genus. Th is fact, which has been noticed in several are used to diff erentiate groups of species. However, genera of Trechini, is also present in other tribes as, for species with either unsclerotized or more or less sclero- example, Harpalini: the genus Parophonus Ganglbauer tized inner sacs can be found in the same genus. Th is is 1892, with one species, Parophonus (Parophonus) iberi- the case of Cimmerites Jeannel 1928, species with pu- parcus Zaballos & García-Múñez 1991 which, unlike bescent integuments and of which the fi rst described the rest of species, lacks sclerotization in the inner sac species showed the inner sac of the aedeagus with- (Zaballos & García-Múñez 1991; Sciaky 1992). out an apparent copulatory piece (Casale & Laney- In the light of above-mentioned data and reasoning, rie 1982). However, Cimmerites species with a more we propose the formal synonymy of Hydrotrechus with or less sclerotized copulatory piece have been recently Trechus, and thus: [Hydrotrechus syn. nov. of Trechus]. discovered (Belousov 1998). Such variety of the endo- A new name for the species must be proposed, as phallus is much more general that what a priori can be it cannot be named “Trechus cantabricus” to avoid thought. Another example can be found in the genus secondary homonymy. Scientifi c names follow the Duvalius, in which most of the species have aedeagus principle of priority (CINZ 2000, art. 23), according with sclerotized isotopic copulatory pieces; however, a to which the valid name of a taxon is the oldest available small number of species lack such structure (Casale & name applied to it. Because a “Trechus cantabricus” Laneyrie 1982, Avon 1997) as, for example, Duvalius Jeannel 1927 already exists, althought it is a synonymy macedonicus (J. Müller 1917) and Duvalius jurjurae of Trechus distigma Kiesenwetter 1851 (Lorenz 1998; (Peyerimhoff 1909). Although for D. macedonicus the Moravec et al. 2003), a substitute name (nomen novum) subgenus Platyduvalius Jeannel 1928 was proposed (CINZ 2000, art. 60) must be proposed. As the junior (and accepted until now; Moravec et al. 2003), it must homonym “cantabricus” has no available synonym, be pointed out that in any moment the peculiarity of it must be substituted by a new name (CINZ 2000,

28 Taxonomic notes on Trechini and a new species art. 60.3). We propose the name Trechus triamicorum example, Trechus, we think it is not appropriate to single nom. nov. as a courtesy to the three (gr. tria) friends A. alberichae and A. leonensis out into Apoduvalius, and (lat. amicorum) who discovered the species (Emilio we propose the synonimization of Trichapoduvalius Carabajal, José García & Fabriciano Rodríguez). with Apoduvalius: Trichapoduvalius syn. nov. of Trechus triamicorum Ortuño & Jiménez-Valverde, Apoduvalius. nom. nov. for Hydrotrechus cantabricus Carabajal, García & Rodríguez 1999 [already occupied by Trechus Habitat and biology of T. cifrianae sp. nov. cantabricus Jeannel 1927, synonym of Trechus distigma Th e only individual of T. cifrianae sp. nov. was Kiesenwetter 1851]. captured inside a cave, approximately 90 m depth, under a stone in a sandbank of the subterranean river The subgenus Trichapoduvalius E. Vives 1976 that crosses the cave. Th e accumulation of organic In the middle of the 20th century, Jeannel (1953) material in this area is scarcer than in the rest of the proposed a new genus in Trechini named Apoduvalius cave and, in consequence, the invertebrate species Jeannel 1953. Th is group of Trechini species is richness and abundance are lower. Among the circumscribed to the hypogean environment of the species that share the habitat with T. cifrianae Cantabrian Cornice (northern Spain) and were assigned sp. nov. we highlight: the troglobite Gastropoda to the “Duvalius phyletic series”, a criterion that is (Pulmonata, Ellobiidae) Zospeum suarezi Gittenberger supported by several authors (Laneyrie 1974; Casale & 1980, an abundant species in other caves of the area Laneyrie 1982) who interpret the arrangement of the and Basque Country (Bellés 1987); the troglophyle copulatory pieces as isotopic. However, Vives (1976) Diplura Campodeidae Campodea pieltaini Silvestri expressed the view that Apoduvalius should better be 1932, widely distributed in the Cantabrian Cornice included in the “Aphaenops phyletic series” because (Sendra & Moreno 2004); two troglobite Coleoptera of the structure of the copulatory pieces of the inner (Leiodidae), Quaestus (Quaesticulus) sharpi nigricans sac (small dimensions and, in his opinion, anisotopic (Jeannel 1924) and Quaestus (Quaesticulus) autumnalis disposition). Th is last criterion is fi nally adopted by autumnalis (Escalera 1898), whose distribution is Casale et al. (1998). Anyway, it is interesting to mention restricted to a few localities in Cantabria (Salgado that, in the Basque-Cantabrian hypogean environment, 1994); several Araneae species (Bellés 1987) as Robertus anophthalmic Trechini (without ommatidia or with a cantabricus Fage 1931 (Th eridiidae), Trichoncus small ocular scar), such as Apoduvalius spp., Trechus pinguis Simon 1926 (Linyphiidae) and Chorizomma pecignai and Trechus triamicorum nom. nov., all show subterraneum Simon 1872 (Dictynidae) and diff erent a clear reduction (even disappearance) of the sclerotized Coleoptera (Carabidae) species (Ortuño & Toribio pieces of the inner sac. 2006) as Iberotrechus bolivari (Jeannel 1913), Trechus After a few years, Español (1971) described a new barnevillei Pandellé 1867, Trechus fulvus Dejean 1831, species of Apoduvalius, A. alberichae Español 1971, Sinechostictus (Sinechostictus) elongatus (Dejean 1831), overlooking the micropubescence of the integument, Ocys (Ocys) harpaloides (Audinet-Serville 1821), a characteristic that, in that moment, was new for the Pterostichus (Pterostichus) cristatus heydenianus Jacobson genus. Th is trait is exposed by Vives (1976), who uses it 1907 and Paranchus albipes (Fabricius 1792). to propose the subgenus Trichapoduvalius Vives 1976. Th e “Cueva del Pis” cave has been the centre of Finally, Salgado & Ortuño (1998) described a second attention of numerous entomologists since, until species with pubescent integument, Apoduvalius recent times (Ortuño & Toribio 2006), it was the only (Trichapoduvalius) leonensis Salgado & Ortuño 1998, known locality for Iberotrechus bolivari Jeannel 1920. which they assign to the subgenus proposed by Vives Th us, it could seem strange that, despite its troglobite (1976). condiction, just only one individual of T. cifrianae Accordingly to the above-mentioned arguments, sp. nov. has been captured after one century of micropubescence should not be used to designate entomological surveys. Probably, T. cifrianae sp. nov. supra-specifi c taxa. Pubescent integument may simply inhabits the Mesovoid Shallow Substratum (MSS) be an adaptation to life in swampy hypogean habitats. of the surroundings or, maybe, the fi ssure network Glabrous species retain micropuntured integument, a (mesocaverns and microcaverns sensu Howarth 1983) trait that, as already mentioned, is closely related to of the cave and that constitutes the “terrestrial-phreatic pubescence. Th us, although micropubescence can be habitat” (sensu Jeannel 1926: 80) of the area. Th is used to identify species, in any way it can be held to would explain the unusual discovery in this well- have phylogenetic information. prospected cave. Since then, new surveys in the cave So, following the same criterion used in, for have not brought new individuals of the species yet.

29 V. M. Ortuño & A. Jiménez-Valverde

Affinities and taxonomic position of T. cifrianae - Pronotum wider than long, margins run regularly sp. nov. curved to pre-angle sinuosity (Fig. 1). Humeral margin highly convex (Fig. 1). Elytra and temples Given the complexity of the internal systematic with pubescence. Bursa copulatrix sacciform, ventro- of the genus Trechus, Jeannel (1927) proposed several lateral folded walls (auricle-shaped) with an anterior “groups of species” to make the taxonomic handling blind prolongation (Figs. 8–9). Length: 4.5 mm easier. However, since then many new species have ...... Trechus cifrianae sp. nov. If we overlook the pubescente of integument, been discovered and a revision of these groups, then accordingly to general body morphology Trechus considering new anatomical criteria (Ortuño & cifrianae sp. nov. is fairly similar to T. pisuenensis. Toribio 2005), is necessary. For example, several species However, female genitalia anatomy (mainly internal (T. pisuenensis and T. triamicorum nom. nov.) of the genitalia) makes these species diff erent (see Ortuño & Basque-Cantabrian region (and so in the geographic Toribio 2005) and suggests that the three species with proximity of T. cifrianae sp. nov.) cannot be easily pubescent elytra are more closely related. Th e similarity included in any established group. Also, T. carrilloi has is especially strong between T. cifrianae sp. nov. and T. no clear systematic position despite being included in carrilloi. the “T. uhagoni-group” (Toribio & Rodríguez 1997). Having said that, and considering that the structure Conclusions of the aedeagus is unknown, in the present study we 1. Micropuncture and micropubescence are are cautious and we do not include T. cifrianae sp. correlated traits (=linked by ontogeny) in Trechus, and, nov. into any established group. However, we will by extension, in Trechini. Th ey should be considered emphasize the most distinctive traits by which T. the same character from a cladistic point of view. cifrianae sp. nov. is clearly distinguished from other 2. Micropubescence is common in endogean and species. Th e pubescence of elytra separates this species troglobite species which are closely related with the from practically all other Iberian Trechus species. “terrestrial-phreatic habitat”. Pubescence probably Th is trait is only present in two other Iberian species enhances survival in swampy environments, retaining which show a close geographic distribution to the air bubbles in immersion conditions and promoting new species, T. carrilloi and T. triamicorum nom. nov. fl otation under sudden fl ooding situations. Several morphological traits separate the new species 3. Neither anophthalmy, micropubescent from these two, as illustrated in the following key: integument nor inner sac of aedeagus without sclerotized pieces, the three traits invoked by Carabajal Iberian Trechus with pubescent elytra et al. (1999), support the existence of the monobasic genus Hydrotrechus. Th e morphological traits of H. 1.- Microphthalmic. Antennae shorter than half the cantabricus Trechus longitude of the body (LRA<0.5) (Fig. 12). Elytra with are in keeping with those of . pubescence. Humeral margin weakly arched. Aedeagus Th e singularity of inner sac must be interpreted as a (Fig. 14) with the median lobe apex upwards hook- synapomorphy, an already known trait in this genus shaped; inner sac with a long sclerotized lamina and as well as in other genus of Trechini (e.g., Duvalius, scaled area. Bursa copulatrix sacciform, with folded among others). walls in the anterior part (auricle-shaped) and with 4. We propose the following synonimization: a short blind prolongation in distal end of bursa (see [Hydrotrechus syn. nov. of Trechus]. Toribio & Rodríguez 1997). Length: 4.1–4.2 mm ...... Trechus carrilloi Toribio & Rodríguez 1997 5. We propose a replacement name (nomen novum) Hydrotrechus cantabricus Trechus triamicorum - Anophthalmic (with small scar). Antennae longer than for : Or- half the longitude of the body (LRA>0.5) (Figs. 10–11) tuño & Jiménez-Valverde, nom. nov. for Hydrotrechus ...... 2 cantabricus Carabajal, García & Rodríguez 1999 [al- 2.- Pronotum cordiform, with lateral margins oblique in ready occupied by Trechus cantabricus Jeannel 1927, basal two thirds (Fig. 15). Humeral margin weakly synonym of Trechus distigma Kiesenwetter 1851]. arched (Fig. 15). Pubescence also present in pronotum 6. Micropubescence should not be used as unique and cephalic disk, but shorter than in elytra and temples. character to designate supraspecifi c taxa. Th us, following Aedeagus (Fig. 13) with the median lobe apex rounded the same criterion as in Trechus, micropubescent (not so acuminated as in the original description); inner sac without sclerotized pieces. Tubular-shaped integuments in Apoduvalius alberichae and A. leonensis bursa with a barely perceptible lateral extension cannot be used to single them out. We propose the (see Carabajal et al. 1999). Length: 3.9–4.3 mm following synonimization: [Trichapoduvalius syn. nov...... Trechus triamicorum nom. nov. of Apoduvalius].

30 Taxonomic notes on Trechini and a new species

7. A new hypogean species of Trechus with Belousov I. A. 1998. Le complexe générique de Nannotrechus Winkler du pubescent elytra is described: Trechus cifrianae sp. nov. Caucase et de la Crimée (Coleoptera, Carabidae, Trechini). Pensoft, Sofi a – Moscow – St. Petersburg, 256 p. Th ree characteristics suggest the Mesovoid Shallow Belousov I. A., Kabak I. 1993. Nouveaux Trechus Clairv. de l’Asie moyenne Substratum (MSS) of the surroundings or the fi ssure (Coleoptera, Carabidae, Trechini). Bulletin de la Société Entomologique network (mesocaverns and microcaverns) of the cave de Mulhouse 1993: 17-29. as the habitat of this species: anophthalmy, pubescent Carabajal E., García J., Rodríguez F. 1999. Descripción de un nuevo integument and the existence of just one known género y una nueva especie de Trechini (Coleoptera: Caraboidea: individual after a century of entomological surveys in Trechidae) de la cordillera Cantábrica. Elytron 13: 123-131. Casale A., Laneyrie R. 1982. Trechodinae et Trechinae du Monde. Tableau the “Cueva del Pis” cave. des sous-familles, tribus, séries phylétiques, genres, et catalogue général 8. Although external morphologies of Trechus des espèces. Memoires de Biospeologie 9: 1-226. cifrianae sp. nov. and T. pisuenensis are fairly similar, Casale A., Vigna Taglianti A., Juberthie Ch. 1998. Coleoptera Carabidae, the structure of internal genitalia space them out and p. 1047-1081 in: Juberthie C., Decu V. (eds.), Encyclopaedia suggests a close relation between the three species with Biospéologica. Vol. 2. Sociéte de Biospéologie, Moulis-Bucarest. CINZ 2000. Código Internacional de Nomenclatura Zoológica. Forth edition. pubescent integuments. Th e main similarity can be [Spain edition]. Consejo Superior de Investigaciones Científi cas- observed between T. cifrianae sp. nov. and T. carrilloi. Museo Nacional de Ciencias Naturales, Madrid, 156 p. 9. Basque-Cantabrian region (Cantabrian Cornice Dethier M., Hubart J-M. 2005. La “troglobitude”: adaptations à la vie and Basque Mountains) shows a high Trechini souterraine. Notes fauniques de Gembloux 57: 29-48. diversity: all known Apoduvalius species (15 sps.), 25 Deuve Th . 1988. Etude phylogénétique des Coléoptères Adephaga: redé- Trechus species (approximately 50% of Iberian species), fi nition de la damille des Harpalidae, sensu novo, et position systé- matique des Pseudomorphinae et Brachinidae. Bulletin de la Société and Iberotrechus bolivari and Hydraphaenops galani Entomologique de France 92: 161-182. Español 1968. Just only 6 Trechus species (T. obtusus, Deuve Th . 1993. L’abdomen et les genitalia des femelles de Coléoptères Ade- T. quadristriatus, T. fulvus, T. schaufussi, T. barnevillei phaga. Mémoires du Muséum National d’Histoire Naturelle 155: 1-184. and T. distigma) show distributions that go beyond Deuve Th . 2002. Deux remarquables Trechinae anophtalmes des cavités the Basque-Cantabrian region, although T. barnevillei souterraines du Guangxi nord-occidental, Chine (Coleoptera, Trechi- and T. distigma have their main distribution in the dae). Bulletin de la Société entomologique de France 107: 515-523. Español F. 1971. Nuevos tréquidos cavernícolas de la fauna española region. Th e other 19 species are Basque-Cantabrian (Col. Caraboidea). Publicaciones del Instituto de Biología Aplicada 51: endemisms. Cantabrian and Basque mountains 89-96. have had major importance in the speciation and Guéorguiev B. V. 2007. Winklerites fodori sp. n. and systematic position of diversifi cation of Trechini, favored by the high diversity Duvalius (Duvalius) fodori Scheibel (Coleoptera, Carabidae, Trechinae). of subterranean, nemoral and orophylic environments Acta Zoologica Academiae Scientiarum Hungaricae 53(2): 107-115. Howarth F. G. 1983. Ecology of cave . Annual Review of (see Serrano 2003). In the last 15 years new Trechus Entomology 28: 365-389. species (T. pecignai, T. carrilloi, T. apoduvalipennis, Israelson G., Palm T. 1979. Th e Trechus species of the Canary Islands T. triamicorum nom. nov., T. pisuenensis, and now T. (Coleoptera: Carabidae). Entomologica Scandinavica 10: 147-151. cifrianae sp. nov.) have been discovered thanks to the Jeannel R. 1926. Faune Cavernicole de la France avec une étude des conditions intensifi cation of surveys of the hypogean environment. d’existence dans le domaine souterrain. Lechevalier, Paris, 334 p. Th ere is no reason to believe that new discoveries will Jeannel R. 1927. Monografi e des Trechinae. Morphologie comparée et distribution géographique d’un groupe de Coléoptères (II livraison). not continue to happen in this region (Ortuño & L’Abeille 33: 1-592. Toribio 2005: 120). Jeannel R. 1936. Les Trechinae des Iles Canaries. Revue française Aknowledgements: We are grateful to Dr. Juan Junoy for d’Entomologie 3(1): 1-18. making available the photographic instrumental. Th e fi rst Jeannel R. 1953. Un genre nouveau de Trechini cavernicoles des monts author wishes to express his eternal gratitude to Dra. Esperanza Cantabriques. Notes Biospéologiques 8: 121-125. Gutiérrez-Redomero for her constant encouragement and Laneyrie R. 1974. Sur la systematique des Trechinae (Coleoptera: friendship. Th e second author was supported by a MEC Trechidae). Nouvelle Revue Française d’Entomologie 4: 3-21. (Ministerio de Educación y Ciencia, Spain) postdoctoral Lorenz W. 1998. Nomina Carabidarum.- A Directory of the Scientifi c fellowship (Ref.: EX-2007-0381). Names of Ground Beetles (Coleoptera “Geadephaga”: Trachypachidae and Carabidae incl. Paussinae, Cicindelinae, Rhysodinae). First edition. References Published by the author, Tutzing, 1190 p. Abeille E. 1903. Description de deux espèces de Trechus aveugles européens Lövei G. L., Sunderland K. D. 1996. Ecology and behavior of ground (Col.). Bulletin de la Société entomologique de France 1903: 298-299. beetles (Coleoptera: Carabidae). Annual Review of Entomology 41: Avon C. 1997. Capitre premier. Evolution et phénomènè de spéciation, p. 231-256. 25-39 in : Avon C. (ed.), Les Duvalius et leur complexe evolutif. Espèces Machado A. 1992. Monografía de los carábidos de las islas Canarias (Insecta, Françaises, Italiennes & Parasites. Edition du L.E.F.H.E., Cagnes Sur Coleoptera). Instituto de Estudios Canarios, La Laguna, 734 p. Mer. Margaría C., Lanteri A. A. 2006. Caracteres taxonómicos. Chapter 4, p. Bellés X. 1987. Fauna cavernícola i intersticial de la península ibérica i les 49-67 in: Lanteri A. A., Cigliano M. M. (eds.), Sistemática Biológica: illes balears. CSIC, Editorial Moll, Mallorca, 207 p. Fundamentos teóricos y ejercitaciones. Ediciones de la UNLP, La Plata.

31 V. M. Ortuño & A. Jiménez-Valverde

Moravec P., Uéno S.-I., Belousov I. A. 2003. Carabidae: Trechinae: Salgado J. M., Ortuño V. M. 1998. Th e new cave-dwelling beetle species Trechini, p. 288-346 in: Löbl I., Smetana A. (eds.), Catalogue of (Coleoptera: Carabidae: Trechinae) of the Catabrian karst (Spain). Th e Palaearctic Coleoptera. Vol. 1. Apollo Books, Stenstrup. Coleopterists Bulletin 52(4): 351-362. Ortuño V. M. 2002. Estado de conocimiento de los artrópodos de Sciaky R. 1992. Revisione dei Selenophorina paleartici occidentali (Coleo- España, p. 209-233 in: Pineda F. D., de Miguel J. M., Casado ptera Carabidae Harpalinae). (XXXVI contributo alla conoscenza dei M. A., Montalvo J. (eds.), La diversidad biológica de España. Pearson Coleoptera Carabidae). Bollettino di Zoologia agraria e di Bachicoltura, Educación, Madrid. Ser. II, 24(1): 37-65. Ortuño V. M., Arillo A. 2005. Description of a new hypogean species of Sendra A., Moreno A. 2004. El subgénero Campodea s. str. en la Península the genus Trechus Clairville, 1806 from eastern Spain and comments Ibérica (Hexapoda: Diplura: Campodeidae). Boletín de la Sociedad Trechus martinezi on the -lineage (Coleoptera: Adephaga: Carabidae). Entomológica Aragonesa 35: 19-38. Journal of Natural History 39(40): 3483-3500. Serrano J. 2003. Catálogo de los Carabidae (Coleoptera) de la Península Ortuño V. M., Outerelo R., Alonso J. 1992. Estudio taxonómico Ibérica. Monografías S.E.A. nº 9. Sociedad Entomológica Aragonesa, comparativo de las especies ibéricas de Chlaeniellus Reitter, 1908 Zaragoza, 226 p. (Coleoptera, Caraboidea, Callistidae). Boletín de la Real Sociedad Toribio M. 1990. Un nuevo Trechus Clairville, 1806 de la provincia de Española de Historia Natural (Sección de Biología) 88: 147-163. Ortuño V. M., Toribio M. 2005. Descripción de un nuevo Trechus Alicante, España (Coleoptera: Trechidae). Elytron 4:181–184. Clairville, 1806 (Coleoptera, Carabidae, Trechini) de los Montes Toribio M. 1992. Un nuevo Trechus Clairville, 1806 del norte de España Cantábricos orientales (norte de España). Graellsia 61(1): 115-121. (Coleoptera: Trechidae). Elytron 6: 87-90. Ortuño V. M., Toribio M. 2006. Ecological relocation of the palaeoendemic Toribio M., Rodríguez F. 1997. Un nuevo Trechus Clairville, 1806 de Iberotrechus bolivari (Jeannel): from troglobiont to epigean (Coleoptera: Cantabria, norte de España (Coleoptera: Carabidae: Trechinae). Carabidae: Trechini). Th e Coleopterists Bulletin 60(1): 23-30. Zapateri, Revista aragonesa de entomología 7: 281-286. Salgado J. M. 1994. Distribución geográfi ca de Speocharis sharpi (Escalera, Vives E. 1976. Coleópteros cavernícolas nuevos o interesantes de la 1898): consideraciones sobre la armadura del saco interno del edeago. Península Ibérica y Baleares. Speleon 22: 159-168. Coexistencia con otros Bathysciinae (Coleoptera, Cholevidae). Nouvelle Zaballos J. P., García-Múñez P. L. 1991. Parophonus iberiparcus sp. n. de Revue d’Entomologie (Nouvelle Série) 11(1): 5-17. carábido ibérico (Col. Caraboidea, Harpalidae). Eos 67: 147-151.

32