taxonID	type	description	language	source
0D1987CFB872FFBAFF654C9DFD3F1C7F.taxon	materials_examined	Type species: Baronniesia delioti Fresneda, Bourdeau & Faille sp. n.	en	Fresneda, Javier, Bourdeau, Charles, Faille, Arnaud (2009): Baronniesia delioti gen. n. sp. n., a new subterranean Leptodirini from the French Pyrenees (Coleoptera: Leiodidae: Cholevinae). Zootaxa 1993: 1-16, DOI: 10.5281/zenodo.185598
0D1987CFB872FFBAFF654C9DFD3F1C7F.taxon	description	Description. The type species of this new genus is characterized by its large size, 4.0 – 5.0 mm long, being one of the largest species of Leptodirini known from the Pyrenean chain. Sexual dimorphism is very accentuated: male with dilated pentamerous protarsi, antennomeres 5 to 9 exceptionally wide, forming a sort of blade (Fig. 1), filiform in females (Fig. 2). Elytra strongly punctured; punctures wide and rough, forming poorly defined transverse striae, especially in the front area. Mesoventral carina low, forming an obtuse angle, and not extended on metaventrite. Male genital segment reduced, forming a slightly sclerotized ring with two very long lateral appendices. Width of aedeagus (Fig. 3) decreasing from base to apex in dorsal view, apex rounded with central part truncated; lateral margins sinuate; aedeagus slightly bent in lateral view (Fig. 4), forming an obtuse angle; terminal tip strongly curved; basal part proportionally large. Lateral styles of aedeagus very thin through their entire length; apex simple, with only three spines (Fig. 7). Inner sac of aedeagus with the three sections typical of the Speonomus phyletic lineage (Fig. 5) (Jeannel 1910, Dupré 1992, Fresneda 1998, Fresneda & Salgado 2000); reinforcement bands (BRA) only in apical area, very long and reaching the base of the section, which is round and more sclerotized, inserted on two additional large feather-like structures located at the external apical part of the FDM (Fig. 5: BDAM). Median area with dorsal (Fig. 5: FDM) and ventral feather-like structures (Fig. 5: FVM), two additional slightly curved feather-like structures forming an inverted “ V ” located above FDM and FVM (Fig. 5: IVP), between BDAM. Apical part of PY in basal area (Fig. 5: FAPY), above connection nodule (NL), very thin and elongate; feather-like structures associated with PY with strongly sclerotized and large basal nodule; ventral basal complex very robust and sclerotized. Eighth female urosternite with short and robust apophyse and round apex (Fig. 15). Spermatheca of “ type 1 ” sensu Perreau (1989) (Fig. 16), thin and elongated, slightly curved and with two well-differentiated lobes; apical lobe spherical and smaller than basal lobe, with spermathecal gland inserted at the same place as spermiduct. Spermiduct 10 – 12 times longer than spermatheca, which gets larger close to the insertion with spermatheca and crosses a sclerotized disc at the contact with bursa copulatrix. Main part of spermiduct close to spermatheca strongly curved.	en	Fresneda, Javier, Bourdeau, Charles, Faille, Arnaud (2009): Baronniesia delioti gen. n. sp. n., a new subterranean Leptodirini from the French Pyrenees (Coleoptera: Leiodidae: Cholevinae). Zootaxa 1993: 1-16, DOI: 10.5281/zenodo.185598
0D1987CFB872FFBAFF654C9DFD3F1C7F.taxon	etymology	Etymology. The name of the new genus refers to the mountainous massif of Baronnies, located north of the central Pyrenees, between the Aure valley and the Adour valley. Taxonomic notes. Baronniesia belongs to the Infraflagellates of the Speonomus “ phyletic series ” of Jeannel (1910). However, the new genus is morphologically very isolated from the rest of the Pyrenean fauna. At first glance, it looks like a large Euryspeonomus breuili Jeannel, 1919 (Sierra de Aralar, Navarra, Spain) but an accurate examination shows that some genital structures are very different, and similar to Antrocharis Abeille de Perrin, 1878 (Ariège, France) and related taxa. The lateral styles of the aedeagus of Baronniesia are very thin through their entire length, with only three spines at the apex. The inner sac of the aedeagus has two additional feather-like structures forming an inverted “ V ” above the FDM and FVM, and the apical part of PY is very thin and elongate. All these characters allow to link the new genus to the Pyrenean genera of Leptodirini of the Antrocharis group (sensu Fresneda et al. 2007), which is redefined below.	en	Fresneda, Javier, Bourdeau, Charles, Faille, Arnaud (2009): Baronniesia delioti gen. n. sp. n., a new subterranean Leptodirini from the French Pyrenees (Coleoptera: Leiodidae: Cholevinae). Zootaxa 1993: 1-16, DOI: 10.5281/zenodo.185598
0D1987CFB872FFBAFF654C9DFD3F1C7F.taxon	description	2) Lateral styles of the aedeagus are shorter than the median lobe, very thin through their entire length and each carrying three long spines at the apex (one shorter), without any penicillum, cavity or membranous lamina (Figs. 8 – 9). The lateral stylus of Baronniesia conforms to this morphological structure (Fig. 7). 3) Presence of two additional strongly sclerotized feather-like structures forming an inverted “ V ”, above the FDM and FVM (Figs. 5 – 6: IVP). These two feather-like structures are present in Baronniesia, which also has two additional structures located laterally of the inverted “ V ” (Fig. 5: BDAM). 4) The apical part of PY (FAPY), above the connection nodule (NL), is very thin and elongate. Baronniesia shares this character with all genera of the Antrocharis group (Fig. 5: FAPY). In all the other genera of the Pyrenean Leptodirini (not members of the Antrocharis group) the apical part of PY is shorter, broader and conical.	en	Fresneda, Javier, Bourdeau, Charles, Faille, Arnaud (2009): Baronniesia delioti gen. n. sp. n., a new subterranean Leptodirini from the French Pyrenees (Coleoptera: Leiodidae: Cholevinae). Zootaxa 1993: 1-16, DOI: 10.5281/zenodo.185598
0D1987CFB872FFBAFF654C9DFD3F1C7F.taxon	description	In all other Pyrenean Leptodirini the number of spines in the apex of the lateral stylus of the aedeagus varies from two to five, but they all have a penicillum, which could be inserted in a depression or not, or be inserted on a membranous lamina. There are two main types of lateral stylus of the aedeagus among the rest of Pyrenean Leptodirini: 1) The Troglophyes type. The apex of the lateral stylus of the aedeagus with two or (more frequently) three long spines, with a dense penicillum formed by numerous short setae. This penicillum may be inserted in a cavity or directly on the integument of the apex of the stylus (Figs. 10 – 12). In the speluncarum group of Speonomus Jeannel, 1908 the third spine is often concealed by the penicillum. The following groups have the lateral stylus of the Troglophyes type: Bathysciella Jeannel, 1906; Bellesia Fresneda & Hernando, 1994; Ceretophyes Fresneda, 1998; speluncarum (Jeannel, 1947) and ehlersi (Perreau, 2000) groups, and the subgenus Machaeroscelis Jeannel, 1924 of Speonomus; Phacomorphus Jeannel, 1908, including the subgenus Phacomorphoides Dupré, 1989 (Fig. 12); Parvospeonomus Bellés & Escolà, 1977 (Fig. 11); Perriniella Jeannel, 1910 and Troglophyes Abeille de Perrin, 1894 (Fig. 10). Troglophyes is a para- or polyphyletic genus (Fresneda et al. 2007). All the easternmost species of Troglophyes, T. bedeli Jeannel, 1906, T. g. gavoyi Abeille de Perrin, 1894, T. g. alluaudi Jeannel, 1911, T. ludovici Chobaut, 1903 and T. oblongulus Reitter, 1908, are closely related to Ceretophyes Fresneda, 1998 and Perriniella Jeannel, 1910, unlike the Antrocharis group of genera and T. a. aubryi and T. a. vallierensis of the central Pyrenees. 2) The Trocharanis type. The apex of the lateral stylus of the aedeagus is with three long spines (sometimes five). One can be longer than the other two (Fig. 13: Speonomus curvipes subcurvipes (Abeille de Perrin, 1878 )), or two longer than the third one, all three of different size (Fig. 14: Paraspeonomus vandeli Coiffait, 1952) or all subequal. A dense penicillum formed by numerous long setae inserted on the side or on a more or less developed membranous lamina is always present; in some species the long setae of the penicillum look like a pedunculus or thin extension of the membranous lamina (e. g. in S. carrerei Fourès, 1954). This group is formed by the genus Paraspeonomus Coiffait, 1952, Speonomus of pyreneus and zophosinus groups (Jeannel 1947), subgenus Metaspeonomus Coiffait, 1959 and Trocharanis Reitter, 1885. All the genera of the southern side of central Pyrenees and coastal ridges of Catalonia (Spain) belong to this group: Lagariella Fresneda, 2000, Naspunius Fresneda, Hernando & Lagar, 1994, Pallaresiella Fresneda, 1998, Salgadoia Fresneda, 1998, Speonomites Jeannel, 1910, Stygiophyes Fresneda, 1998, Trapezodirus Jeannel, 1924 and Troglocharinus Reitter, 1908 (except the three taxa of the subgenus Antrocharidius Jeannel, 1910 with a secondary loss of penicillum but with the membranous lamina). Dilated antennae in male of Baronniesia is a character shared with Josettekia angelinae Bellés & Déliot, 1983 and Paraspeonomus vandeli Coiffait, 1952. Josettekia angelinae has the 9 th (especially) and 10 th male antennomeres dilated. However, in Josettekia this is not a generic character, as the second species of the genus, J. mendizabali (Bolívar, 1921), does not show any dilatation of the male antennae. Both species of Josettekia have the apex of the lateral stylus with four short spines and lack the two inverted “ V ” feather-like structures in the inner sac of aedeagus. In P. vandeli only the fourth antennal article of male is dilated, but the lateral stylus of the aedeagus has a membranous lamina and a penicillum, and the inner sac lacks the two inverted “ V ” feather-like structures. Phylogenetic analyses. Baronniesia gen. n. and the genus Gesciella (omitted from the previous analyses) were added to the data matrix of Fresneda et al. 2007 (Appendix 1: this paper) with the aim of including both genera in a formal phylogenetic analysis. See Fresneda et al. 2007: 335 – 344 for the list of the 34 characters and their states. For a list of studied species see Appendix 1 in Fresneda et al. 2007. For composition of groups of terminal taxa sharing identical character states and lumped together for the purposes of analysis see Appendix 2 in this paper. Despite the fact that three new characters were added to the matrix of Fresneda et al. (2007) (see below), this could not help to resolve relationships between Baronniesia and the most closely related taxa (the Antrocharis group). In the new character matrix (Appendix 1) Baronniesia shares all character states with Antrocharis, Gesciella, Paratroglophyes and Troglophyes. Character 35. Lateral styles of aedeagus: very thin along their entire length (0) and always shorter than the median lobe (Figs. 3 – 4); or strong along their entire length (1), with the apex more or less thickened and of length ranging from shorter to longer than the median lobe. Coded as missing data (?) for the species with character states 26 (1), 27 (1), 28 (0), 29 (0), 30 (1) and 31 (1), sensu Fresneda et al. (2007). These species have the apex of the lateral stylus of the aedeagus with two to five spines; when they have three, there is always a penicillum, cavity or membranous lamina. Character 36. Apical part of the median region of the inner sac of aedeagus: with two very sclerotized pieces forming an inverted “ V ” located above FDM and FVM (Fig. 5: IVP) (1); without inverted “ V ”, or with other structures (0). In the species with the inner sac of the aedeagus lacking copulating armature (Fresneda et al. 2007: character 16, state 0) character 36 was coded as missing data (?). Character 37. Basal region of inner sac of aedeagus: apical piece of PY (FAPY) above the connection nodule (NL) very thin and elongated (Fig. 5: FAPY) (1) or short and conical (0). In species without PY (Fresneda et al. 2007: character 17, state 0) character 37 was coded as missing data (?). The extended data matrix was analysed using the same methods as in Fresneda et al. (2007): the shortest trees were heuristically searched in PAUP 4.0 b 10 (Swofford 2002) with 10,000 tree-bisection-reconnection (TBR) replicas, swapping on all multiple starting trees, and saving all of them. To obtain a higher resolution, data were reweighted successively according to the rescaled consistency index (Swofford, 2002), and a heuristic search conducted on the initial set of the shortest trees (see Fresneda et al. 2007 for details). The topology of the strict consensus of the 330 reweighted trees (consistency index 0.67, retention index 0.9) was identical to that of Fresneda et al. (2007) (Fig. 17) and, thus, was not affected by addition of three new characters. Gesciella and Baronniesia appear in the same polytomy with the remaining genera of the Antrocharis group, as they share all character states.	en	Fresneda, Javier, Bourdeau, Charles, Faille, Arnaud (2009): Baronniesia delioti gen. n. sp. n., a new subterranean Leptodirini from the French Pyrenees (Coleoptera: Leiodidae: Cholevinae). Zootaxa 1993: 1-16, DOI: 10.5281/zenodo.185598
0D1987CFB874FFB1FF654DBDFD0C1B44.taxon	materials_examined	Type locality. France, Hautes-Pyrénées, Esparros, subterranean river of Artigaléou-Arodets. Type series. Holotype: FRANCE: ɗ, Esparros (massif of Baronnies), subterranean river of Artigaléou- Arodets, 10. V. 2007, Bourdeau & Faille leg. (Muséum National d’Histoire Naturelle, Paris). Paratypes: 33 ɗɗ and 61 ΨΨ, same locality, data and collectors (authors’ collections). Holotype description. Habitus and size. Baronniesia delioti sp. n. is one of the largest species of Pyrenean Leptodirini. Length from anterior edge of pronotum to apex of elytra: 4.54 mm (see Variability). Anophthalmous and depigmented; outline oval, stocky, with pronotum wider than elytra (Fig. 1). All the surface covered by yellowish pubescence, thin and soft; elytral pubescence longer. Punctation of head thin; mandibles very robust with a subapical tooth at internal edge. Antennae. Length 3.50 mm; antennal articles gradually dilated from fourth to ninth, tenth and eleventh articles not dilated; ventral side of the ninth article concave, dorsal side strongly convex. For measurements of antennal articles see Table 1.	en	Fresneda, Javier, Bourdeau, Charles, Faille, Arnaud (2009): Baronniesia delioti gen. n. sp. n., a new subterranean Leptodirini from the French Pyrenees (Coleoptera: Leiodidae: Cholevinae). Zootaxa 1993: 1-16, DOI: 10.5281/zenodo.185598
0D1987CFB874FFB1FF654DBDFD0C1B44.taxon	description	Pronotum. Strongly transverse (length 1.25 mm, width 2.30 mm), sides very arcuate, maximum width before posterior edge. Elytra. One and half time longer than wide, strongly convex, with dehiscent apices; each elytron regularly curved at apex. Punctation strong and rough, in basal area arranged in transverse striae; in apical half with punctures distributed irregularly. Legs. Proportionally long; mesotibiae arched, metatibiae straight. Male protarsomeres 1 – 3 strongly dilated, the first wider than apex of protibia; onychium as long as protarsomeres 1 – 4 combined. Mesoventral carina, aedeagus and male genital segment as in the description of the genus. Female diagnosis. Habitus as in Fig. 2; body smaller than in male (see Variability), pronotum as wide as elytra; outline perfectly oval. Antennae filiform, articles not dilated. For measurements of antennal articles see Table 1. Protarsi tetramerous, non dilated. Urosternite VIII and spermathecal complex as in the description of the genus. Variability. Body length (measured from anterior edge of pronotum to apex of elytra): male: 4.02 – 4.54 mm, average 4.30 mm; female 4.01 – 4.40 mm; average 4.16 mm.	en	Fresneda, Javier, Bourdeau, Charles, Faille, Arnaud (2009): Baronniesia delioti gen. n. sp. n., a new subterranean Leptodirini from the French Pyrenees (Coleoptera: Leiodidae: Cholevinae). Zootaxa 1993: 1-16, DOI: 10.5281/zenodo.185598
0D1987CFB874FFB1FF654DBDFD0C1B44.taxon	etymology	Etymology. This new species is dedicated to our entomologist colleague and friend Philippe Déliot, who died recently.	en	Fresneda, Javier, Bourdeau, Charles, Faille, Arnaud (2009): Baronniesia delioti gen. n. sp. n., a new subterranean Leptodirini from the French Pyrenees (Coleoptera: Leiodidae: Cholevinae). Zootaxa 1993: 1-16, DOI: 10.5281/zenodo.185598
0D1987CFB874FFB1FF654DBDFD0C1B44.taxon	distribution	Distribution and ecology. The new species is only known from a single cave located in the central part of the Pyrenean chain, south of Esparros. The total length of the cave of Artigaléou is more than one kilometer, with two levels of chalk-stoned fossil galleries before reaching the subterranean river. Specimens of B. delioti were found only in traps placed near the river. Apparently, the species does not occur in the deeper parts of the cave, and it seems likely that the specimens found were carried from deep soil by water together with Speonomus bastideus Coiffait, 1950 a and the species of Trechini mentioned below. Hypogean fauna of the Baronnies has never been studied in detail, and there are scarce faunistic data from only three other caves (Jeannel 1928, Coiffait 1950 a). To better characterize the ecology of Baronniesia it would be necessary to investigate other localities where it could live, probably as an MSS or endogean species together with Geotrechus discontignyi, a typical endogeous ground beetle also found near the traps. Biogeographical notes. In the area where B. delioti was discovered, the only other known Leptodirini are the species of the speluncarum group of the genus Speonomus, distributed between the Lez valley (Ariège) and the western Pyrenees, where they have also colonized the southern range in Navarra until the Echo valley (Huesca, Spain).	en	Fresneda, Javier, Bourdeau, Charles, Faille, Arnaud (2009): Baronniesia delioti gen. n. sp. n., a new subterranean Leptodirini from the French Pyrenees (Coleoptera: Leiodidae: Cholevinae). Zootaxa 1993: 1-16, DOI: 10.5281/zenodo.185598
0D1987CFB874FFB1FF654DBDFD0C1B44.taxon	description	The pattern of distribution of the Pyrenean Trechini is presented in Fig. 18. The species of the subgenus Aphaenops Bonvouloir, 1862 have radiated in the western part of the Pyrenean chain, and their diversity increases from Lourdes to the west, the majority of them (including the endogean Geaphaenops Cabidoche, 1965) diversified between Lourdes and Saint-Jean-Pied-de-Port (Faille 2006). The eastern-most French species of the subgenus Aphaenops is A. leschenaulti Bonvouloir, 1862, known from the caves around Bagnères-de-Bigorre. On the contrary, the species of the subgenus Cerbaphaenops Coiffait, 1962 have radiated in the eastern part of the Pyrenees between Ariège valley and Bigorre area, and their diversity decreases from east to west (Faille 2006). The surrounding area of Bagnères-de-Bigorre is the contact area between the two main radiations of subterranean cave Trechini, Aphaenops in the narrow sense in the west and Cerbaphaenops in the central-eastern Pyrenees. In this area, the two lineages are represented by A. (Cerbaphaenops) crypticola ssp. aeacus Saulcy, 1864 and A. (Aphaenops) leschenaulti Saulcy, 1864, both sympatric in the cavities around Bagnères-de-Bigorre, in particular the Castelmouly and Tuco caves (Jeannel 1928). While the species of the subgenus Aphaenops on the northern slope of the range do not reach the area east of Bagnères-de-Bigorre, on the southern slope some species of this subgenus are distributed further east, till the Ribagorçana valley, where A. catalonicus Escolà & Canció, 1983 is present. The pattern of diversification of the Pyrenean Leptodirini is quite similar, although with some differences (Fig. 19). The species of Speonomus of the speluncarum group are distributed from the western Pyrenees to the Aure valley (Sarrancolin, Lortet) and the Labastide area (Artigaléou, Labastide cave). Members of the speluncarum group are found again in Ariège, with S. orgibetensis Gers, 1989 (speluncarum group sensu Gers, 1989: Fig. 1 D), the two species of the ehlersi group, S. ehlersi Abeille de Perrin, 1872 and S. opisthonoxus Gers & Dupuis, 1988 (Gers & Dupuis 1988: Fig. 3 C), and the subgenus Machaeroscelis. The distribution area of the speluncarum group and related Leptodirini are thus extended to the east. The presence of other genera apparently related to the speluncarum group (Ceretophyes, Parvospeonomus, Perriniella, and Troglophyes) in the eastern part of the Pyrenees is very remarkable. A revision of the north Pyrenean Leptodirini would be necessary to elucidate this biogeographical enigma. When Leptodirini are compared to troglobitic Trechini, the main difference in their distribution is that in Leptodirini the contact area of the two groups, the western one (Speonomus of speluncarum and ehlersi groups, Machaeroscelis, Bathysciella and Phacomorphus) and the central-eastern one (Paraspeonomus, Trocharanis and Speonomus of the pyreneus group) is extended to the east until Ariège. The area between Ariège and Bigorre seems to be an overlapping area of distribution for the Leptodirini of the Trocharanis type and those of the Speonomus of the speluncarum group. The lineage of the Antrocharis group of genera seems to have originated in this contact area, but we do not know whether from the western or the eastern group. The coincidence of the same contact area for the eastern and western Pyrenean groups of Leptodirini and Trechini suggests the same geologic and climatic causes.	en	Fresneda, Javier, Bourdeau, Charles, Faille, Arnaud (2009): Baronniesia delioti gen. n. sp. n., a new subterranean Leptodirini from the French Pyrenees (Coleoptera: Leiodidae: Cholevinae). Zootaxa 1993: 1-16, DOI: 10.5281/zenodo.185598
