THE GENUS BATHYSCIOLA JEANNEL , 1910 IN THE IBERIAN PENINSULA AND PYRENEES . TAXONOMIC REVISION OF SECTIONS IV , VI AND VII ( JEANNEL , 1924 ) ( COLEOPTERA , CHOLEVIDAE , LEPTODIRINAE )

The taxonomy of the Iberian and Pyrenean species of Bathysciola Jeannel, 1910 (sections IV, VI and VII of Jeannel, 1924) is revised. The proposed ordenation is based on the study of the genital structures of both sexes, in particular the sclerotised shield of the internal sac of the aedeagus. According to the different models of internal sac were identified the following species groups in the genus Bathysciola Jeannel, section IV: aubei group of Peyerimhoff (1 taxon): B. aubei (Kiesenwetter); madoni new group (2 taxa): B. madoni Jeannel and B. penicillata Jeannel; zariquieyi new group (2 taxa): B. zariquieyi zariquieyi Bolívar and B. zariquieyi serratensis Coiffait; section VII or ovata group of Perreau (8 taxa): B. asperula asperula (Fairmaire), B. asperula subasperata (Saulcy) (= intermedia Jeannel), B. ovata ovata (Kiesenwetter), B. ovata aragonica Coiffait, B. ovata catalana Coiffait, B. ovata gabasensis Hustache, B. simonis (Abeille de Perrin) and B. talpa (Normand); section VI: lapidicola new group (4 taxa): B. arcuatipes Jeannel, B. lapidicola lapidicola (Saulcy), B. lapidicola rectipes Coiffait and B. lapidicola simplex Coiffait; meridionalis new group (3 taxa): B. finismillennii sp. n., B. grenieri (Saulcy) and B. meridionalis (J. Du Val) (= nitidula Normand); larcennei new group (2 taxa): B. convena Jeannel new rank and B. larcennei (Abeille de Perrin); schiodtei group of Perreau (9 taxa): B. bigerrica Jeannel new rank (= convexa Coiffait n. syn.), B. breuili Bolívar (= azuai Bolívar n. syn.), B. diegoi Salgado & Fresneda, B. fauveli Jeannel, B. grandis (Fairmaire), B. obermaieri Bolívar, B. parallela (Jeannel), B. rugosa (Sharp) and B. schiodtei (Kiesenwetter) (= navarica Coiffait). Incertae sedis: Bathysciola aranensis Coiffait and Bathysciola minuscula (Abeille de Perrin).


Introduction
divided the genus Bathysciola Jeannel 1910 into eight sections.The ordenation proposed by this author is based on the geographical distribution and morphological characters, so that species with different models of masculine genitalia, spermathecal complex and occulate or anophthalmic, are grouped together.The only precedent is that Jeannel (1924a) formed his section IV based on the "Bathyscia the aubei group of Peyerimhoff (1905)".The ordenation proposed by Jeannel (1924a) is still valid; the only change made concerns the naming of the sections, which has been replaced by "group species" in the manner of "groupe schiodtei" or other proposals made by Perreau (2000).This genus, which now contains 83 species distributed from the western Pyrenees to the Caucasus could be compared to a sack containing taxa of dubious phylogenetic value.The need for a review of the complex has already been suggested by Vailati (1988) who, with regard to the "Bathysciola complex", states that "... ancora resta in attesa di migliore definizione"; also Giachino (1998) points out that the systematic position of the genus is dubious and is currently being completely restructured.Fresneda & Salgado (2000) state that the genus Bathysciola appears in several dichotomies of the identification key proposed for Iberian Leptodirinae: this is an extremely heterogenous group which "will probably have to be divided into different genera".Of the sections proposed by Jeannel (1924a) only the fourth, sixth and seventh have Iberian species.The fourth section includes the type species of the genus so this taxon will be taken as the genus model; however, this section includes a number of species distributed in the Provence area (France and Italy) which have not been studied as the aim of this paper is to attempt to order Iberian fauna on both sides of the Pyrenees.The species in this section not found in the Iberian Peninsula will be studied at a later date.The species in the sixth and seventh sections have all been studied, as they are distributed preferably in the Pyrenees; only one species in the sixth section inhabits areas at a considerable distance from the Pyrenees and it has also been studied.The other sections of the genus (sensu Jeannel, 1924a) will be considered some time in the future; the species in these sections are also absolutely heterogenous as regards models of genitalia: the entire genus should perhaps be fragmented into several genera for reasons of coherence.

Material and methods
The following described species and subspecies have been studied: B. aranensis Coiffait, B. arcuatipes Jeannel, B. asperula asperula (Fairmaire), B. asperula subasperata (Saulcy) (= intermedia Jeannel), B. aubei (Kiesenwetter), B. azuai Bolívar, nación tiene como base el examen de las estructuras genitales de los dos sexos, y de un modo especial el estudio de la armadura esclerotizada del saco interno del edeago.Se han diferenciado los siguientes grupos de especies en el género Bathysciola Jeannel, section IV: grupo aubei de Peyerimhoff (1 taxon): B. aubei (Kiesenwetter); madoni nuevo grupo (2 táxones): B. madoni Jeannel y B. penicillata Jeannel; zariquieyi nuevo grupo (2 táxones): B. zariquieyi zariquieyi Bolívar y B. zariquieyi serratensis Coiffait; sección VII o grupo ovata de Perreau (8 táxones): B. asperula asperula (Fairmaire), B. asperula subasperata (Saulcy) (= intermedia Jeannel), B. ovata ovata (Kiesenwetter) The aim of this study is to characterize the different models of internal sac of the aedeagus and to group together those species corresponding to this criterion to provide data for a future ordenation; however, a new generic category has not been established as this should be done as part of a revision of the whole genus.The characterised models are determined by different sclerotized pieces, their shape and position in the sac: support and copulatory pieces (Dupré, 1992).Other genital characters, such as the number of setae on the tip of the lateral style of the aedeagus or the median lobe itself, have been added to this criterion.There is no doubt about the validity of using the internal sac of the aedeagus to form natural groups and this as been used by several authors since Jeannel (1910), who used these characters in the first reordenation of "Silphides cavernicoles".Jeannel (1911) insists that these structures could be the basis for systematic ordenations; therefore, they enable the phylogenetic history of the species in question to be restructured.He states that each species has a consistent and specific copulatory shield corresponding to «... en quelque sorte à la signature d'une série phylétique» (Jeannel, 1922).Jeannel (1924a) published his revision of the Bathysciinae Horn 1880, giving a general description of the different models of internal sac on which he based his proposed ordenation.Jeannel (1950) proposed the ordenation of the genus Speonomus Jeannel 1908, based on the tip of the lateral style.Finally, Jeannel (1955) established what is considered to be the most primitive division of the Bathysciinae: "supraflagellates" and "infraflagellates".A decade later Laneyrie (1967Laneyrie ( , 1969Laneyrie ( and 1978) ) proposed a new ordenation based on the copulatory shield of the internal sac, basically the structures in the basal region.Guéorguiev (1974aGuéorguiev ( , 1974bGuéorguiev ( and 1976) ) proposed a new ordenation using the external morphology: the series of combs and spurs on the tibiae.He established dichotomies for four genera only, based on the parts of the aedeagus.Bellés et al. (1978) proposed a new ordenation for the Iberian species using studies by Jeannel, Laneyrie and Guéorguiev, and they basically use the characters of the male genital structures (internal sac, tip of the lateral style, etc) to include new records on Iberian fauna.Bellés (1983) first used the characters of the internal sac of the aedeagus to establish a dichotomy between Speonomus puncticollis Jeannel 1910and S. espinosai Bellés 1983. Bellés (1984) carried out a detailed study of the internal sacs of two species in the genus Parvospeonomus Bellés & Escolà 1977 and also established the naming of their different pieces.In studies by Bellés, or later, Bellés & Comas (1986), the need for a revision of the species of Iberian Leptodirinae is underlined in the light of a detailed study of the characters of the internal sac of the aedeagus.With these criteria Comas & Escolà (1989) described the subgenus Ceretophyes Comas &Escolà 1989, andComas (1983) reviewed the genus Anillochlamys Jeannel 1909.Following the same line Dupré (1989Dupré ( , 1990Dupré ( and 1991) ) reviewed and ordered the Leptodirinae from the western confines of the Pyrenees until, in 1992, he presented an ordenation based on a detailed and very complete study of the internal sac of the aedeagus; he also redefined the divisions proposed by the previously-mentioned researchers (Dupré, 1992).Fresneda & Hernando (1994) and Fresneda et al. (1994) used the same method to describe the genus Bellesia Fresneda & Hernando 1994 and the subgenus Naspunius Fresneda, Hernando & Lagar 1994. Salgado (1975, 1993, 2000) ordered the Iberian supraflagellates in the series Quaestus Schaufuss 1861 and Speonomidius Jeannel 1924 in groups of species with the same model of internal sac of the aedeagus; Fresneda (1998) and Fresneda & Salgado (2000) did something similar with the Iberian "infraflagellates".Also in the same line is the latest revision of the series Spelaeochlamys (= section Anillochlamys) by Salgado & Fresneda (2003).The latter studies (Salgado, 2000, Fresneda, 1998and Fresneda & Salgado, 2000) also consider some elements of the female genitalia -in the line of Perreau (1989)-adding observations on characters that had not previously been studied: detailed structure of the spermathecal complex (length and thickness of the spermathecal duct, type of insertion in the sper-matheca and in the bursa copulatrix); this is an open line of investigation with few available data, but which appears to be quite promising.In summary, in the last hundred years numerous authors have proposed various ordenations based on different characters to reflect the phylogeny of a group; these contributions have been complementary in some cases and contradictory in others, but all of them make the debate more interesting.The constant increase in described taxa underlines the need to search for new characters or reinterpret traditionally used ones.Following this line of work, Bellés (1984) consolidated the basis of the study of the sclerotised shield of the internal sac of the aedeagus; it has also been followed by different authors including Fresneda et al. (in press) who demonstrated that these are the most consistent characters when establishing phylogenetic relationships based on a binary matrix with multiple characters according to the cladistic method used.In any case, this paper does not include a phylogenetic study to establish relationships between the different groups of species; its aim is to define groups that will provide a reasonably simple identification of the species.In a later study, which has already been started, an effort will be made to establish relationships between these taxa and other genera of Leptodirinae inhabiting the same regions.
All the insects in this study should be included in the Leptodirinae infraflagellates.Contributions by Jeannel (1924aJeannel ( , 1955)), Laneyrie (1967), Guéorguiev (1974a, b and1976), Casale et al. (1991) and Giachino et al. (1998) are considered, but the meaning given to the term infraflagellates is that contributed by Dupré (1992) who redefines this group based on the study of histological sections of the internal sac of the aedeagus: models of internal sac with a Y-shaped piece in the basal region, and dorsal phanerae in the median region.The spermathecal complex in all these genera correspond to "type 1" of the spermatheca defined by Perreau (1989).
The terminology used to designate the different pieces of the copulatory shield of the internal sac of the adeagus according to the denominations established by Bellés (1984), Bellés & Comas (1986), Dupré (1992) and Fresneda (1998).For each model the situation and name of the structures considered are indicated in the corresponding illustration.Whenever necessary, a new name is included if there is no equivalent piece in the quoted referente (see Figs. 1,5,12,15,31,42,48,80).
As for the specific identity of the taxa of a determined group, all those species with differences in the genital structures have been considered as different species, and those with identical genitalia but different external morphology as subspecies.These criteria are argueable, however an effort has been made to apply an aseptic system to give some coherence to the proposed ordenation and mimimize as far as possible the observer's own perception.There are several subspecies described and in most cases the description is based on a small number of specimens, often only one; an effort has been made not to directly establish synonymies with the respective nominotypic subspecies as the person responsable for the description did not consider the constancy of the differential characters in question in a population; this is also the reason why there are not enough data to deny the validity of these taxa; all the subspecific names in the same situation have been kept in the hope that future samplings will shed some light.
Section IV of Jeannel, 1924Bathysciola Sectio IV of Jeannel, 1924. Arch. Zool. exp. gén., 63(1): 104 The Bathysciola species lacking transverse striolae on the elytra but with a sutural stria not parallel to the suture were grouped together by Jeannel (Jeannel, 1924a) and named section IV.This section includes the true Bathysciola as it includes the type species of the genus; in the Iberian Peninsula there are three species, belonging to two groups.In his check list Perreau (2000) establishes the correspondence between "Bathysciola Section IV Jeannel, 1924" and"Groupe aubei of Peyerimhoff, 1905"; in this study the two categories are maintained because although B. madoni, B. penicillata and B. zariquieyi do, in fact, belong to section IV, they cannot be included in the same group as B. aubei.
GEOGRAPHICAL DISTRIBUTION.The species in the madoni group are found in the northern peninsula area, from the north of the western Pyrenees to the pre-Pyrenean mountains in the south, in interior regions of Catalonia; there are not many data on captures and their distribution areas are marked by a number of collection points at a considerable distance from each other.Jeannel, 1923Bathysciola madoni Jeannel, 1923. Bull. Soc. entomol. Fr., 28: 104 Bathysciola (Bathysciola) madoni Jeannel, 1923: Jeannel, 1924. Arch. Zool. exp. gén., 63(1): 109 Bathysciola (Bathysciola) madoni Jeannel: Perreau, 2000.
Je.The other three specimens are females.Two of them are considered syntypes.The first because it has the same collection inscription as the male, "Tourau"; and the second because it has "feuilles" written on it, and the original description states: "... , en tamisant des feuilles, ..." (Jeannel, 1923).The third one is not included in the type series because it bears the word "piquets", a term that does not appear in the description by Jeannel. First Puigsacalm, 22.VII.1951, Montada leg. (Lagar, 1954;Español, 1956); S-B, Montesquiu, X.1934, Español leg. (Lagar, 1954;Español, 1956); S-GI, Camprodón (Español, 1956).The following specimens are in the col.Coiffait preserved at the MNHNP: S-GI, Ripoll, 28.XII.1955, HC leg., 1 ex.;19.V.1956, HC leg., 4 exs. (Coiffait, 1959); S-GI, Ribes de Fresser, 2.XI.1955, HC leg., 1 ex.;26.XII.1955, HC leg., 4 exs.;5.IV.1956, HC leg., 12 exs. (Coiffait, 1959).MALE GENITALIA.Median lobe of aedeagus in dorsal view with mid region expanded (Fig. 5); in lateral view (Fig. 6) uniformly arcuate from one end to the other and of uniform thickness, even the apical third.The tip of the lateral style of the male of the type series (Fig. 7) bearing twelve arcuate setae of similar length.Internal sac (Fig. 5) with symmetrical branches of Y-shaped piece prolonged over sides of the sac to median region where they come into contact with the appendage of the dorsal sacs; strongly sclerotized U-shaped piece where the axial piece originates; there is a sac densely covered in fine pilosity amongst the reinforcing bands, and another one covered in small thick spines between the hyaline protruberances of the Y-shaped piece; V-shaped piece amongst reinforcing bands.
DISCUSSION.The male genitalia of a specimen from Prats de Mollo in France was illustrated.With regard to B. aubei, the differences are enormous, especially in the internal sac of the aedeagus; the genital structures of these two taxa cannot be considered as belonging to the same model (Figs. 1 and  5).B. madoni is extremely similar to B. penicillata.The difference between these species is not very clear.The specimens collected in the area of Ripollès (Gerona) were identified as B. penicillata by Jeannel (preserved in the MNHNP general collection) and B. madoni by Coiffait (Coiffait, 1959); the latter author points out that the number of setae on the tip of the lateral style of the aedeagus varies considerably (eight to fourteen), which was perhaps the main character justifying the description of B. penicillata; it was also discovered that the lateral style preserved from the single type specimen of B. penicillata has ten setae in the distal extreme, and not nine as indicated in the description.The internal sac of both species is very similar, but the median lobe is quite different in lateral view: uniformly arcuate in B. madoni (Fig. 6) and subrectilinear with the apical third sinuous in B. penicillata (Fig. 9).The determination of all the Iberian references of this species should be reviewed as it is highly likely that many specimens classified as B. madoni are in fact B. penicillata.The genitalia of the male of the type series was extracted from the glass slide cover and prepared again on an acetate paper slide placed in Canada Balsam and inserted on the same pin as the specimen it belongs to; the two lateral styles had been separated from the median lobe by R. Jeannel.
symetrical branches of the Y-shaped piece prolonged along the sides of the sac to median region where they come into contact with the appendix of the dorsal sacs; strongly sclerotized U-shaped piece where the axial piece begins; sac densely covered in fine pilosity between the reinforcing bands and another covered in thick small spines between the hyaline prolongations of the Y-shaped piece.FEMALE GENITALIA.Spermatheca (Fig. 11) curved throughout, both lobes much larger than median region.DISCUSSION.Jeannel (1924b) described this species from one specimen.The genitalia of this specimen was prepared between two slides; it is completely flattened laterally and one of the lateral style is separated; it was prepared again on an acetate paper slide in Canada Balsam and inserted on the same pin as the specimen it belongs to.Only the tip of the lateral styles of this specimen was illustrated (Fig. 10) as the rest is completely deformed; the remaining structures weren illustrated using the type locality specimens.The number of setae on the lateral style of the aedeagus varies considerably and overlaps with the number of setae in B. madoni.Only the shape of the median lobe of the aedeagus enables the correct determination, which has already been indicated in the discussion on B. madoni.
The zariquieyi new group DIAGNOSIS.This group is characterised by the internal sac of the aedeagus (Fig. 12) with evident ventral phanerae (Fig. 12: VPM), and very small dorsal ones (Fig. 12: DPM) prolonged until coming into contact with the Y-shaped piece (Fig. 12: YP); apical reinforcing bands (Fig. 12: ARB) resting upon membranous ligulae (Fig. 12: ML) whose tegument is densely covered in spines; these ligulae have a sclerotized peduncle in the inferior region.The tip of the lateral style of the aedeagus armed with 5-6 setae.Only one species with two subspecies.The term «group" is used though we are dealing with only one species in coherence with the proposed ordenation.
GEOGRAPHICAL DISTRIBUTION.Distributed in mountains situated on the Catalonian coast.
MALE GENITALIA.In lateral view, median lobe of aedeagus (Fig. 13) arcuate in basal two thirds, which are more or less thick, in apical third lobe subrectilinear or sinuous and decreasing uniformly to tip; there may be a small depression on the dorsal face; distal extreme pointed or peak-shaped.Lateral style fine, slender, shorter or as long as median lobe and with tip inflated in the shape of an elyptic club bearing five setae, two longer and three shorter (Fig. 14).Internal sac (Fig. 12) with symetrical branches of the Y-shaped piece prolonged along sides of the sac to median region where they come into contact with a quadrangular sclerotized piece; two large sacs covered in spines at base of apical region.
DISCUSSION.The internal sac of the aedeagus of one specimen of the type series was extracted and the two lateral styles separated; it is prepared between two glass slides, and separated from the specimen it belongs to.In the illustrations, the internal sac of the aedeagus and tip of the lateral style belong to this specimen, whereas the other structures were illustrated from a specimen from the Montseny massif.The original description (Bolívar, 1919) indicates that the tip of the lateral style of the aedeagus has five setae, two longer and three shorter ones; the datum is correct for the two lateral styles of the studied male of the type series, but Jeannel (1924a, b) states that the specimens he studied have six.This taxon is quite isolated; the internal sac of its aedeagus is very different to that of the other Iberian species in section IV.It should probably have its own genus, but this division should be made from the perspective of a revision of the genus, especially the species in section IV.For the moment, no taxonomical changes are to be made.
Section VII of Jeannel, 1924 or ovata group of Perreau, 2000Bathysciola Sectio VII of Jeannel, 1924. Arch. Zool. exp. gén., 63(1): 113 Groupe ovata of Perreau, 2000. Mém. de la SEF, 4: 230 DIAGNOSIS.The Bathysciola in section VII are characterized by not having a parasutural stria, and the transverse striolae on the elytra.The internal sac of the aedeagus (Fig. 15) has an axial piece (Fig. 15: AP) and very small phanerae (Fig. 15: PM) joined together in the median region, which may be triangular or practically forming a blunt nodule (Figs. 19,22 and 25); the apical reinforcing bands (Fig. 15: ARB) rest on membranous ligulae (Fig. 15: ML) whose tegument may be covered in large spines.The internal sac of this group of species is very similar to that of Iberian species in the madoni group.Four species.
GEOGRAPHICAL DISTRIBUTION.Both sides of the Pyrenees, French Massif Central and western foothills of the Alps, near Lyon (France).(Kiesenwetter, 1850) MALE GENITALIA.In lateral view (Fig. 16) median lobe of aedeagus with basal two thirds rectilinear and base expanded; apical third folded in obtuse angle, strongly attenuated and sinuous; lateral style (Fig. 17) fine, sinuous and shorter than median lobe; bearing three long setae of similar length at tip, one of which possibly slightly shorter and finer; upper part of median region of internal sac of aedeagus (Fig. 15) with two sclerotized bands covered in long, fine spines; amongst them the shield of the axial piece is formed by two pieces joined together in a U-shape bearing a fine stylet in the middle; just below there are two triangular phanerae with an expanded interior margin also bearing a number of strongly robust spines.FEMALE GENITALIA.Spermatheca (Fig. 18) short, rectilinear and robust; two differentiated spherical lobes joined by a cylindrical median region; spermathecal duct very short, as long as spermatheca; small nodule at insertion point and sclerotized tubular piece at point of contact with bursa copulatrix.Ovipositor corresponding to the model described for the phyletic series Speonomus (Fresneda, 1998); nevertheless, articulated distal piece not triangular but very fine with base slightly expanded at articulation point.

Bathysciola (Bathysciola) (ovata group) ovata ovata
DISCUSSION.The original description does not indicate the number of specimens studied and the specimen or the specimens of the type series have not been found in the investigations carried out in different museums.So, apart from the interpretation of the original description (Kiesenwetter, 1850), the characterisation of the species provided by different authors who state that they have identified the species (Bolívar, 1919;Jeannel 1911Jeannel & 1924a;;Español, 1956;Coiffait, 1959) has been accepted.Numerous specimens of French origin determined by R. Jeannel and H. Coiffat deposited in the M NHNP collection, have been located and studied.The original description indicates the type locality: "Bagnières de Luchon" (Kisenwetter, 1850); different samplings carried out in the locality did not provide any specimens though some were found in a nearby locality; alter comparing them with the MNHNP specimens and verifying that it was the same species, they were used for the descriptions and illustrations in this study.A number of specimens from Artiga de Lin, Baix Aran, Lérida (Spain) were used for the description and illustrations.Bathysciola ovata belongs to section VII of the genus Bathysciola proposed by Jeannel (1924a).Considering the genital characters the relationship between B. ovata and the Iberian Bathysciola in section IV (B.madoni and B. penicillata) is quite evident: the internal sac of the aedeagus has a number of sacs with the tegument covered in spines, very short, triangular ventral phanerae and a robust sclerotized shield on the axial piece resembling a nodule situated in the upper part of the median region of the sac.The external morphology cannot be confused with other species and genera due to the peculiar facies: very small and apparently spherical.There is another character which is not found in the other specimens in the series of Iberian Speonomus or in the other three species in the group: the mesosternal carina is prolonged over the metasternum in a similar way to most of the specimens in the series Quaestus (supraflagellates).Five subspecies which are difficult to place were described.One of them (Bathysciola ovata minuscula Abeille de Perrin, 1901) has been considered in the section incertae sedis as the single specimen of the type series is a female and its identity and relationship have not been determined.The complex is distributed on both sides of the Pyrenees.
DISCUSSION.A series of specimens from the Grotte de l'Herm, Ariége (France) was used in the description.These specimens are identical to the specimen of the type series found in the col.MNHNP.The male genitalia of Bathysciola asperula is different to that of B. ovata, including the median lobe and in particular, the internal sac.It is similar to B. simonis and related to B. talpa.The most similar species is B. simonis.The internal sacs of the aedagus of these two taxa are similar but the median lobe is shaped differently: it is uniformly curved in B. simonis, the basal portion being much more robust, whereas the basal portion in B. asperula is rectilinear and elongate.Also, the lateral styles -in lateral view-are fine and slender in B. asperula and more robust in B. simonis.As for B. talpa: lateral style fine to tip and the distal tip of the fine, weakly curved median lobe in B. asperula are very different to the thick lateral style ending in a club and the thick strongly curved tip of the median lobe in B. talpa; in any case although the relationship between these taxa must be distant, their genitalia have a vaguely similar facies.B. asperula is distributed in the French departments of Ariége and Aude on the northern slope of the Pyrenees.(Jeannel, 1909)  COMPARATIVE STUDY.The male genitalia is identical to that of the nominotypic subspecies; they are basically differentiated from the latter by being slightly larger.Secq & Secq (1996) established the synonym B. asperula intermedia with B. asperula subasperata.The lectotype and paralectotypes of these taxa were designated in the same study.In the original description of Adelops subasperatus (Saulcy, 1872) the number of studied specimens was not specified.

Bathysciola (Bathysciola) (ovata group) simonis
(Abeille de Perrin, 1875) In the original description of B. simonis the number of studied specimens is not indicated: "Découvert par M. Eugène Simon dans des mousses, au Lioran (Cantal)" (Abeille de Perrin, 1875).The specimens of the type series were not found.The following specimens from the MNHNP general collection were studied: F-Tarn et Garonne, Lacaune, alt.850 m., Galibert leg., 4 males and 5 females; F, Le Lioran, 28.10.1892,Grouvelle leg., 1 female; F, Le Lioran, Falcoz leg., 1 male; F, Aveyron, La Cavalerie, 17.7.1925, 1 female.MALE GENITALIA.Median lobe of aedeagus slender (Fig. 23) and uniformly curved from base to apical third; in lateral view, from this point sinuous, pointed tip slightly expanded and curved; lateral style almost as long as median lobe; robust, arcuate with distal region strongly folded towards median lobe; bearing three subequal setae inserted in distal extreme (Fig. 24); internal sac of aedeagus (Fig. 22) with few weakly sclerotized pieces; Yshaped piece in basal region with fine arms; median region with two elongate nodules touching on the axial line of the sac, set out on a triangular piece and covered in striate tissue; reinforcing bands without any outstanding characters.DISCUSSION.A series of specimens from Lacaune in the department of Tarn (France) were used for the description and illustrations.The most similar species is B. asperula, which is clearly related.The internal sacs of the aedeagus of these taxa are very similar but the shape of the median lobe is quite different: rectilinear and elongate in B. asperula and uniformly arcuate in B. simonis, in the latter species the basal portion is much more robust.The lateral styles, in lateral view, are more robust in B. simonis than in the other species.This taxon was described as a species but considered by Jeannel (1910) as a subspecies of B. asperula.ver, differences in the genitalia of the two taxa are enormous: the very thick lateral style ending in a club and the thick strongly curved tip of the median lobe in B. talpa contrasts heavily with the lateral style which are fine to the tip, and distal tip of the fine, weakly curved median lobe in B. asperula.The six strong spines in the median region of the internal sac of B. talpa are found exclusively in this species.The first segment of the male protarsi in B. talpa is also remarkable: it is much wider than the distal extreme of the protibia and heart-shaped.There is no similar case amongst the Leptodirinae in this study in which the first segment is always equal, slightly wider or narrower than the protibia.This species was considered a subspecies of B. asperula by Jeannel (1910) and bona species by Coiffait (1959); Normand (1907) posed the possibility of describing it as a subspecies subordinate to B. asperula.
The lapidicola new group DIAGNOSIS.The male genitalia of Bathysciola lapidicola (Saulcy, 1872) is taken as the model.A series of specimens from Aubert, Ariège (France) were used for the description and illustrations.This group is characterised by the more or less slender aedeagus in lateral view (Figs. 29 and 32), and folded in an obtuse angle in the mid region; there may be an expanded ventral area in the upper part of the basal region; in dorsal view it is robust, with almost parallel sides, weakly converging to the slightly truncate tip; lateral style somewhat shorter than median lobe and four tiny setae in line transversally inserted in tip (Figs. 30 and 33); internal sac of aedeagus (Figs.28 and 31) characterized by two elongate, triangular shaped phaneroid complexes (Fig. 31: PCM), situated on either side of the symmetry axis of the sac, in the median region; the apical reinforcing bands (Fig. 31: ARB) rest upon membranous ligulae (Fig. 31: ML).This complex is related to the Bathysciola of the meridionalis group because of certain similarities in the internal sac of the aedagus: it is as though the basal pieces of the two longitudinal phaneroid complexes of the lapi-dicola group were joined by their internal margin in the same way as the phanerae in the meridionalis group, and forming a U-shape or almost a ring formed by several pieces.There is no similarity with other models of internal sac.In any case the differences between the internal sac of lapidicola and the meridionalis group are considerable and affect the number, shape and position of the sclerotized shield.Two species.
GEOGRAPHICAL DISTRIBUTION.The group is distributed throughout low altitude mountainous regions in the French departments of Haute Garonne and Ariège, on the northern slope of the Pyrenees.

Bathysciola (Bathysciola) (lapidicola group)
arcuatipes Jeannel, 1924 Bathysciola (Bathysciola)  There are another four specimens labelled but without the «TYPE" label, in the MNHNP general collection.Finally, one last specimen is included in the description of the species: Cazavet / 21.IV.1914/ Dodero.It should be pointed out that whilst the description of the taxon is dated 1924, the type series -as indicated by the label handwritten by Jeannelwas collected in 1926 by Jeannel himself; however, there is no doubt about its identity as one of the specimens had the "TYPE" red label.R. Jeannel may have made a mistake when labelling or the number corresponding to 1924 of the Arch.Zool.exp.gén.may have been published late.The following specimens are also found in the MNHNP general collection: F-Ar., Aubert, Grotte d'Aubert, Argod leg., 2 exs.Also in the MNHNP, but in the Coiffait collection are: F-Ar., Aubert, Grotte d' Aubert, 18.6.1943, HC leg., 3 exs.;28.9.1945, HC leg., 2 exs.; F-Ar., Foret de Lestelas, VII.1958, HC leg., 1 ex.MALE GENITALIA.Aedeagus in lateral view (Fig. 29) uniformly curved and robust, especially the basal portion, without expanded ventral area in the median region; tip prolonged and very pointed.Lateral style tip with four tiny setae transversally set out in a line (Fig. 30).Internal sac of aedeagus (Fig. 28) characterised by two elongate triangularshaped phaneroid complexes, situated on both sides of the symmetry axis, in median region; upper area of these pieces prolonged in bundles of weakly defined twisted fibres in the upper parts of the internal sac; the two lateral phanerae of tip of basal region (floating pieces of other models of internal sac, Fresneda, 1998) large and noticeably sclerotized to upper extreme.
DISCUSSION.The possibility of transferring B. arcuatipes to the B. lapidicola complex as another subspecies has been discussed.However, not enough specimens have been studied, particularly those in the subspecies B. lapidicola; they are only known from the type localities and only the specimens in the type series, normally one or two specimens.The existing specific and subspecific table is to be provisionally maintained until a good representation of all the taxa is obtained.This species is closely related to B. lapidicola simplex, as they have similar male metatibiae: triangular and slightly curved in the apical fourth.The basal region of the aedeagus of B. arcuatipes (Fig. 29) is more robust and does not have a ventrally expanded area which is characteristic of B. lapidicola (Fig. 32); their respective internal sacs of the aedeagus are very similar and it is difficult to define small differences between B. arcuatipes (Fig. 28) and B. lapidicola (Fig. 31).Jeannel (1924a) indicates that the type series is from Grotte de Lestelas; but the located specimens are labelled as being from Aubert, where they must have been found under stones.MALE GENITALIA.In lateral view (Fig. 32) aedeagus slender and folded in obtuse angle in the middle more or less; ventrally expanded area in upper part of basal region; in dorsal view, robust, with almost paralell sides, weakly converging in slightly truncate tip; lateral style somewhat shorter than median lobe, four tiny setae transversally set out inserted in tip (Fig. 33); internal sac of aedeagus (Fig. 31) characterised by two elongate, triangular phaneroid complexes on either side of the axis of symmetry of the sac, in median region; the upper part of these pieces is prolonged in a kind of well-defined translucid phanera in the upper part of the internal sac; the two lateral phanerae of the tip of the basal region (floating pieces of other models of internal sac, Fresneda, 1998) are small, upper extreme weakly sclerotised.
DISCUSSION.Specimens in the type series from Aubert, Ariége (France) were used to illustrate the male genitalia.Bathysciola lapidicola is closely related to B. arcuatipes: see discussion on B. arcuatipes.Bathysciola lapidicola is a complex of four subspecies based on inconsistent characters.In the subspecies described, there is some variation in the shape of the median lobe of the aedeagus and the internal sac; there are also some insignificant differences in secondary sexual characters, such as the shape of the male metatibiae.Two of the subspecies have been described based on a very small number of specimens-one and two specimens, respectivelyand the differential characters have such little significance that the option of establishing the synonymy with the nominotypic subspecies has been considered; it was decided to preserve them provisionally until new specimens were collected: if the differential characters, though small, exist in all the population and if the population has a determined distribution area, the category of subspecies would be justified.

Bathysciola (Bathysciola) (lapidicola group)
lapidicola rectipes Coiffait, 1959  COMPARATIVE STUDY.In lateral view aedeagus tends to be slightly more robust than type form but with ventrally expanded area in upper extreme of basal region.It is differentiated from lapidicola by the straight, slender metatibiae, expanded from the first third and then subparallel to the tip.The number of specimens in the type series is not indicated and a male specimen is described."Type: Gouffre Djerbao vers 1.100 m à Arbas, Haute-Garonne (M.Bouillon).Ma collection."(Coiffait, 1959).This specimen is preserved in the col.robust than in the nominotypic subespecies and has a ventral expanded area.It is differentiated from lapidicola by the triangular, slightly curved metatibiae in the apical fourth; also, by the first segment of the male protarsi which is as wide as the distal extreme of the protibia, slightly narrower in lapidicola.
The meridionalis new group DIAGNOSIS.The male genitalia of Bathysciola meridionalis (Jacquelin du Val, 1854) was taken as the model.Median lobe of aedeagus (Fig. 43) curved just beyond central area, forming obtuse angle; apical part slightly shorter; four short setae inserted two by two (Fig. 44) in tip of lateral style; internal sac (Fig. 42) with a series of strongly sclerotised phanerae in median region (Fig. 42: PCM), an upper one and a smaller lower one, both tranversal, the ends of which are joined by a fairly sclerotised and strongly wrinkled tissue; in the central part, in axial position, strongly sclerotised floating nodule (Fig. 42: FN); inferior part of apical reinforcing bands (Fig. 42: ARB) with a sac (Fig. 42: SS), the tegument of which is strongly striate, with very sclerotised robust spines on the upper part.Spermathecal complex similar to B. finismillennii n. sp.(Fig. 37).Spermatheca curved from one end to the other and both lobes differentiated, their section being larger than that of the median region; spermathecal duct inserted in spermatheca via a nodule; expanded, striate and crossing a kind of sclerotised cylinder at the point where it joins the bursa copulatrix; spermathecal gland connected to terminal nodule of spermathecal duct, at the point where it joins the spermatheca.On evaluating the characters of the internal sac of the aedeagus, no similarity with any other model was found.It is true that the tip of the style has four setae, as in the Bathysciola of the schiodtei group, or Euryspeonomus Jeannel, 1919or Speocharidius Jeannel, 1919; however, this is not a conclusive character in establishing relationships as there is also a species in this group with nine setae whose internal sac leaves no doubt of the taxonomic position of the meridionalis group.Three species.GEOGRAPHICAL DISTRIBUTION.Western half of the northern slope of the Pyrenees, occasionally, some localities on the southern slope; also, the Garona river basin, as far as Bordeaux.
DESCRIPTION.Body length: 2,30 mm.Oblong, oval-shaped and uniform.Surface covered in fine, yellowish, laid back pilosity.Pronotum transverse, with lateral margins uniformly arcuate and tegument marked with clear, dense disordered punctures with a dull appearance.Elytra lacking parasutural striae and sculpture formed by rugose points vaguely lined up transversally.Mesosternal carina low and uniformly arcuate.Legs proportionally short and first segment of male protarsi quadrangular, as wide as distal extreme of protibia.MALE GENITALIA.Lobe of aedeagus (Fig. 35) strongly arcuate in central region, almost forming a right angle and the two resulting halves practically of the same length; nine long setae (Fig. 36) inserted in tip of lateral lateral style; median region of internal sac (Fig. 34) with a series of strongly sclerotized phanerae forming anular complex; in the centre, in axial position, there is a strongly sclerotized elongate nodule apparently floating; in the inferior part of the apical reinforcing bands there is a sac with a striate tegument.FEMALE GENITALIA.Spermatheca (Fig. 37) uniformly curved from one end to the other and the two lobes differentiated, though their section is much larger than that of the median region; spermathecal duct inserted in the spermatheca via a nodule, and also expanded, striate and crossing a sclerotized cylinder at the point where it joins the bursa copulatrix.
DISCUSSION.All the specimens in this group have a uniformly curved median lobe from one end to the other in lateral view, that is, the median curvature is large and the two resulting parts form a very obtuse angle, the distal part being smaller than the basal one (Figs. 35,39 and 43); four setae of varying length -short (Fig 44) or long and twisted (Figs 36 and 40)-depending on the species, inserted in the apex of the lateral lateral lateral style.In B. finismillennii n. sp. the median lobe is strongly curved in the central region almost forming a right angle, and the two resulting halves are practically of the same length (Fig. 35); nine long setae inserted in the tip of the lateral style (Fig. 36).B. meridionalis is the most similar of all the species in the group.It is practically impossible to differentiate it from this species by the external morphology and the internal sac of the aedeagus is also very similar; with a strongly sclerotized nodule in axial position in the upper part of the median region and a sac with a striate tegument.However, it can be differentiated from this species and others, as previously commented, by the supranummary setae on the tip of the lateral style of the aedeagus.The presence of numerous setae on the tip of the lateral style has already been described for other Iberian specimens in the genus Bathysciola: B. zariquieyi (with five or six setae), B. madoni (with eight to fourteen setae) and B. penicillata (with nine or ten setae).In any case these taxa are at a considerable distance from the species described because the model of internal sac of the aedeagus is completely different, and they are related to the Bathysciola in the aubei group.The specimens in the type series were found in the Coiffait collection (MNHNP) where they had been determined by this author as Bathysciola meridionalis.In the general collection of this museum there is another female specimen (Grotte d'Eycheil, 25.V.1897, J. Bepmale leg.) which is not included in the type series because as it is a female it cannot be determined with absolute certainty; however this specimen was presumibly studied by R. Jeannel in his revision (Jeannel, 1911) and he attributed it with doubts to Bathyscia nitidula; he states that he observed some differences in this taxon in the antennae and suggests the possibility of it being a new species.Years later in the monograph, no comment was made about including the specimen in the studied Bathysciola meridionalis (= Bathyscia nitidula).
ETYMOLOGY.This compound name is formed by the latin terms: finis (end), mille (thousand) and annus (year).The genitive indicates that it is the Bathysciola from the end of the millenium, as this species was discovered during the year two thousand on the Gregorian calendar.

Bathysciola (Bathysciola) (meridionalis group)
grenieri (Saulcy, 1872)  Coll. A. Argod. 1931;5. (hw) grenieri;6. (i, hw, in red): Paralectotype male or female, Bathysciola grenieri (Saulcy), B. Secq dés.1994.This species was recorded on one occasion from Andorra, without specifying the locality (Español, 1956).MALE GENITALIA.A series of specimens from Ax les Thermes, Ariége (France) was used for the description and illustrations.Base of aedeagus in lateral view (Fig. 39) robust and strongly folded in middle, apical half decreasing brusquely becoming very fine, tip prolonged and pointed or ending in small nodule; lateral style sinuous, as long as or slightly shorter than median lobe with tip expanded and club-shaped with nine (Fig. 36) or four setae of varying length (Figs. 40 and 44); in dorsal view internal sac of aedeagus (Fig. 38) characterised by strongly sclerotized complicated phanera superposed in median region; occasionally forming a type of ring with an interior space ocuppied by a floating nodule; upper extreme of phanerae and between reinforcing bands of apical region, there may be one or several sacs with a strongly striate tegument; in lateral view phanerae in median region forming a kind of parenthesis.
DISCUSSION.This species is easily differentiated within its group as its relationship with B. finismillenni n. sp. is as distant or as close as with B. meridionalis.The latter two species are a lot more similar to each other.Bathysciola grenieri and the other species have the same model of internal sac when observed in lateral view as they both have bracket-shaped phanerae, however, in dorsal view B. grenieri does not have strongly striate sacs, and the floating nodule inside the sclerotized ring in the other species is modified in this one, and formed by three phanerae joined by one end, resembling the letter T. The description by Saulcy does not specify the number of specimens studied.Lectotype and paralectotypes were erroneously designated in the study by Secq & Secq (1996).The lectotype and paralectotypes in fact correspond to Adelops grenieri, not to Bathysciola grenieri as indicated on the labels included by Secq and as indicated in the mentioned study.MALE GENITALIA.The genital structures of B. meridionalis were chosen as the model for charac-terising the group: internal sac of the aedeagus (Fig. 42), aedeagus in lateral view (Fig. 43) and tip of the lateral style (Fig. 44).
DISCUSSION.The type series comes from Bordeaux.The male genitalia was illustrated using a series of specimens from Monferran, in the department of Gers, France.Of the species in the group the most similar one is B. finismillennii sp.n.The internal sac of the aedagus of these two taxa has the same layout and appearance: with a strongly sclerotized nodule in axial position in the upper part of the median region and a large sac with a strongly striate tegument.However, there  meridionalis.42) internal sac of the aedeagus in dorsal view.43) aedeagus in lateral view.44) apical region of the lateral style.Scale bar: a-b, 0.5 mm: 42, 43.ARB, apical reinforcement band; FN, floating nodule; PCM, phanerae complex of the medial region; SS, striated sac; YP, Y-shaped piece.are easily discernible differences between the respective sacs, as can be seen in figures 42 (B.meridionalis) and 34 (B.finismillennii sp.n.).The shape of the median lobe is also different, it is longer and more angulose in B. finismillennii sp.n., and there are nine setae inserted in the tip of the lateral style of this species compared with four in B. meridionalis.
The larcennei new group DIAGNOSIS.The male genitalia of the species Bathysciola larcennei (Abeille de Perrin, 1883) is taken as the model.A series of specimens from Pordiac in Gers, France was used in the description and illustrations.In lateral view (Fig. 49) median lobe of aedeagus strongly expanded ventrally in basal region and folded in uniform curve just before half its length; distal half gradually decreasing and tip pointed and slightly curved ventrally.Lateral style slightly shorter than median lobe, bearing four long subequal setae and a membranous lamina on tip (Fig. 50).Upper area of median region of internal sac of aedeagus (Fig. 48) with two small phanerae joined together by a small sclerotized bridge (Fig. 48: PM); in lower region and at a distance from these phanerae there is a transversal lamina (Fig. 48: TL) bearing two fine long superposed phanerae set out obliquely; inferior region of these phanerae with two identical nodules (Fig. 48: SN) with a peduncle prolonged to the area near the Y-shaped piece (Fig. 48: YP).Facies of internal sac of aedeagusnumber, shape and layout of sclerotized shieldshow no great similarity with any of the other models described.Apart from the differences there is some similarity with the model of the schiodtei group in the layout of the groups of phanerae: two situated at both ends of the median region.As in the meridionalis or schiodtei groups the tip of the lateral style has four setae which also occurs in the Basque-Navarre genera Euryspeonomus or Speocharidius; as previously indicated, this is not a determining character when establishing degrees of relationship if it is not as a complement of other more conclusive ones.The need to study the other genera of Leptodirinae inhabiting these territories, is underlined.Two species.GEOGRAPHICAL DISTRIBUTION.Northern slope of the central Pyrenees, in the departments of Gers, Haute Garonne and Hautes Pyrénées (France).

Bathysciola (Bathysciola) (larcennei group) convena
Jeannel, 1924 new rank Bathysciola (Bathysciola) larcennei subsp.convena Jeannel, 1924. Arch. Zool. exp. gén., 63 (1): 92 Bathysciola larcennei subsp.convena Jeann.: Coiffait, 1952.Notes biospéol., 7: 44 Bathysciola larcennei convena Jeannel, 1924: Secq & Secq, 1996.l'Entomologiste, 52 (1): 12 Bathysciola (Bathysciola) larcennei convena Jeannel: Perreau, 2000.Mém. de la SEF, 4: 237 MATERIAL STUDIED.The number of specimens is not indicated in the original description.Lectotype and paralectotypes are designated in the study by Secq & Secq (1996).Lectotype male labelled: 1. (hw): Grotte de Gargas; 2. (hw): H. Pyr., R. J., 944,17.VIII.19;3. (i,in red): Type; 4. (i): Muséum Paris, Coll.R. Jeannel. 1931;5. (i, hw): Lectotype male, Bathysciola larcennei convena Jeannel, B. Secq dés.1994.Paralectotypes one male and two females labelled: 1. (hw): Grotte de Gargas; 2. (hw): H. Pyr., R. J., 944, 17.VIII.19;3. (i or hw, in red): Type; 4. (i or hw): Muséum Paris, Coll.R. Jeannel. 1931;5. (i, hw): Paralectotype male or female, Bathysciola larcennei convena Jeannel, B. Secq dés. 1994.MALE GENITALIA.In lateral view (Fig 46) basal region of median lobe weakly expanded ventrally and folded in uniform curve just before half its length; distal region decreasing gradually with dorsus somewhat depressed; tip pointed and slightly curved in ventral part.Lateral style a little shorter than median lobe, with four long twisted setae of similar length at tip (Fig 47).In the upper part of the median region of the internal sac of the aedeagus (Fig 45) there are two small pilous strongly sclerotized phanerae; in the inferior region and at a distance from them there are dorsal and ventral phanerae of peculiar shape, formed by a number of twisted laminas.DISCUSSION.A series of specimens from the Grotte de Gargas, Ariège (France) was used for the description of the male genitalia.This taxon was described (Jeannel, 1924a) as a subspecies of Bathysciola larcennei, and its status had not been modified.The differences observed amongst their respective genital structures justifiy the independance of both species.The median lobe of the aedeagus in lateral view is similar in both species, however a closer study reveals some subtle differences: in B. convena (Fig. 46) the basal region is thick but has no peculiar expanded areas and the distal half decreases gradually with the dorsus slightly depressed; however, in B. larcennei there is a prominent rounded lump in the ventral area (Fig. 49).The lateral style of B. convena (Fig. 47) do not have a membranous lamina at the tip, but B. larcennei (Fig. 50) does.The internal sac of the aedeagus though of the same model in both species, has very different pieces: the upper part of the median region in B. convena has two small pilous strongly sclerotized phanerae whereas in B. larcennei there are two smooth phanerae formed by a number of lamina joined together by a small sclerotized bridge; the differences in all the pieces of the sclerotized shield are numerous (Figs. 45 and 48).MALE GENITALIA.-Thegenital structures of B. larcennei were used to characterise the group; aedeagus in dorsal view and internal sac by transparency (Fig. 48), aedeagus in lateral view (Fig. 49) and tip of lateral style (Fig. 50).DISCUSSION.It is differentiated from the other species in the genus in the discussion on B. convena.In the description by Abeille de Perrin the number of specimens studied is not indicated.The lectotype and paralectotypes were erroneously designated in the study by Secq & Secq (1996).This series of specimens corresponds, in fact, to Bathyscia larcennei and not to Bathysciola larcennei as indicated by the labels that Secq & Secq (1996) included and as stated in the study carried out by these authors.The schiodtei group of Perreau, 2000 Groupe schioedtei of Perreau, 2000. Mém. de la SEF, 4: 236 DIAGNOSIS.The male genitalia of the Bathysciola schiodtei (Kiesenwetter, 1850) was taken as the model.A series of specimens from Vielha, Lérida (Spain) was used for the description and illustrations.For the spermathecal complex, specimens of B. rugosa from Dolina de Orobe in Alsásua, Navarra (Spain).The median lobe in lateral view (Fig. 81) is long, weakly arquate, uniformly decreasing from the base to the tip and with a sinuous dorsal face; the lateral style are as long as the median lobe, pointed and bearing four setae (Fig. 82); the internal sac of the aedeagus (Fig. 80) divided into three areas: in the basal region the Yshaped piece (Fig. 80: YP) formed by two very fine symmetrical rods with a thick bean-shaped nodule articulated by a tubule with another nodule of the same characteristics, at the tip, there are phanerae next to the Y-shaped piece (Fig. 80: NPYP); in the median region ventral (Fig. 80: VPM) and dorsal (Fig. 80: DPM) phanerae weakly sclerotized, not easily discernible; a number of well-defined phanerae visible on both sides of the sac (Fig. 80: LPM); at the end of the median region there are some strongly sclerotized nodules articulated amongst each other (Fig. 80: PC); in the apical region reinforcing bands (Fig. 80: ARB) resting upon sclerotized nodules (Fig. 80: SN).Spermatheca (Fig. 79) curved from one end to the other and median region very thin; both lobes differentiated, the basal one being much larger and more elongate than the apical one: the spermathecal duct is long and fine, and expanded near the bursa copulatrix where it crosses a sclerotized disk; a spermathecal gland can be seen.The schiodtei group is related to the genera inhabiting the Basque-Navarre region, Euryspeonomus and Speocharidius having in common the tips of the lateral style of the aedeagus armed with four differentiated setae and phanerae next to the Y-shaped piece in the basal region of the internal sac.The differences amongst the described taxa in this group are clear and consistent for all the studied specimens; they are all considered valid as they are clearly characterised.However, the differences are not extremely important and the problem arises when assigning a taxonomical category to them: species or subspecies.There are two possible solutions: subordinate all the taxa as subpecies of the oldest taxon or consider all of them as good species.Data in entomological literature demonstrate that the Leptodirinae species do not have very wide distribution areas: the observations made by the authors of this study are along the same line.So the second option has been chosed, that is, to consider practically all the described taxa as species.
GEOGRAPHICAL DISTRIBUTION.The specimens in this group inhabit the Pyrenean mountains, in particular both slopes of the western half, where they diversified into numerous taxa.They also inhabit the northern areas, following the course of the river Garona to its mouth and there is a considerably isolated population in the region of Calvados, northern France.The group probably invaded the Pyrenees going from west to east as phylogenetically similar genera: Euryspeonomus and Speocharidius are found in the western extreme Basque-Navarre territory.It is the only genus of the series that has moved east, perhaps due to being less developed and therefore less capable of colonizing any cryptic habitat, such as forests area that probably stretched uninterrupted between these regions and between both slopes of the Pyrenees.
MATERIAL STUDIED.Subsp.bigerrica: the original description only states: "Type: un mâle de Aspeigt" (Jeannel, 1930).This specimen is stored in the MNHNP general collection and labelled: 1. (i): Ossau / Aspeigt; 2. (hw): Hustache; 3. (i): TYPE; 4. (i): MUSÉUM PARIS / Coll.R. Jeannel 1931.Subsp.convexa: the number of specimens is not indicated in the original description, only: "Type dans ma collection" (Coiffait, 1959).This specimen is stored in the Coiffait collection at the MNHNP and labelled: 1.: (i) B. Pyrénées / (hw Coiffait) LARRAU / (i) H. Coiffait / (hw Coiffait) 22.9.49; 2. (hw Coiffait): Grotte / Aker leccia; 3. (hw Coiffait, in red): TYPE.MALE GENITALIA.In lateral view (Fig 52) median lobe long, weakly arcuate and uniformly decreasing from base, which is very thick to tip with slightly concave face just before; styl lateral style us almost as long as median lobe, pointed and bearing four strongly folded setae similar in length (Fig. 53); in the internal sac of the aedeagus (Fig. 51) there are some phanerae next to the weakly sclerotized Y-shaped piece; in the median region the ventral phanerae are well sclerotized and appear to form the interior margin of translucent laminas; the dorsal ones are hyaline; there are a number of short phanerae can be seen along the sides of the sac, which are not strongly sclerotized and rest upon the laminas of the ventral phanerae; the basal nodules of the phaneroid complex are folded in an acute angle; the interior margin of the reinforcing bands is noticeably expanded, sclerotized and prolonged forming peduncles; amongst the reinforcing bands there is a tiny tubule with a fine rod DISCUSSION.Bathysciola bigerrica was described (Jeannel, 1930) as a subspecies of B. parallela.The differences observed in the latter species have led to this taxon being considered a species for questions of coherence with the criterion applied in this revision.It is similar to a number of species in its own group that have a clearly defined structure between the reinforcing bands of the internal sac of the aedeagus, which is either a tubule, a rod or both; these species are:   Bolívar, 1921 Bathysciola (Bathysciola) breuili Bolívar, 1921. Bol. Soc. esp. Hist. nat., T. 50º aniv.: 525 Bathysciola (Bathysciola) schiodtei subsp.breuili Bolívar, 1921: Jeannel, 1924. Arch. Zool. exp. gén., 63(1): 110 Bathysciola (Bathysciola) schioedtei breuili Bolívar: Perreau, 2000. Mém. de la SEF, 4: 238 Bathysciola (Bathysciola) azuai Bolívar, 1921. Bol. Soc. esp.Hist.nat., T. 50º aniv.: 524 MATERIAL STUDIED.Subsp.breuili: in the original description Bolívar (1921), the author states that he collected and studied various specimens collected by H. Breuil and R. Jeannel on 18th August, 1919; he also states that the "Tipo: macho…" is in col.Biospeologica (in the MNHNP) and the remaining specimens divided between this museum and the MNCNM.The following species belonging to the type series were found: one specimen stored in the MNHNP general collection and labelled:: 1. (hw) 1. 945 cuev.de Landarbaso; 2. Guipuzcoa RJ 3. Bathysciola breuili C. Bol.;4. (i,in Bolívar (1921) the author claims to have collected and studied three specimens which have been located in the museums indicated by these author.One male ("Tipo: macho…".Bolívar, 1921) Mairuelegorreta, 2000, JF and Salgado leg., 16 exs. stored in col. JF and col. Salgado; there is another specimen in the MNCNM collection: S-SS, Rentería, Cueva de Landarbaso, VIII.1923,Abajo leg., 1 male.Another labelled specimen from the MNHNP has been studied: Peña de Gorbea, 1000 m, feuilles, IX.1919, R. Jeannel.MALE GENITALIA.Median lobe of aedeagus in lateral view (Fig. 55) curved in mid region and preapically bisinuate; style with four long subequal setae at tip set out in twos and intricately twisted (Figs.56 and 57); internal sac (Fig. 54) with very fine, weakly sclerotized exterior phanerae; basal phanerae of phaneroid complex strongly folded in obtuse angle; reinforcing bands expanded in basal region forming laminas almost touching in the axis of symmetry of sac.FEMALE GENITALIA.Spermathecal complex coinciding with description given for the genus, spermatheca (Fig. 58) proportionally short and robust, with two noticeably expanded lobes.DISCUSSION.Specimens from the Cueva de Landarbaso, in Rentería (Guipúzcoa) were used in the description of the genitalia.This species was described as a species (Bolívar, 1921) and shortly afterwards (Jeannel, 1924a) was included as subspecies of B. schiodtei.Nevertheless, the exclusive characters of the internal sac of the aedeagus, and a curious secondary sexual character not seen in the genus, justify its status as valid species.This taxon should be included in the schiodtei group which has no differentiated structure amongst the reinforcing bands; these species are: B. breuili, B. fauveli, B. rugosa and B. schiodtei.The most similar amongst them is B. rugosa as they have very similar basal nodules in the phaneroid complex: in an acute angle in B. breuili and obtuse in B. rugosa.In contrast, the other two species in the subgroup have rectilinear nodules with an nocked inferior margin.The lateral phanerae of the internal sac are robust and strongly sclerotized in B. rugosa, whereas in B. breuili they are hyaline and finer; the phanerae adjacent to the Y-shaped piece are very evident in B. breuili but in B. rugosa they are almost invisible.Also, in B. breuili there is a femoral spine in the male metafemurs, not present in B. rugosa or the other taxa in the genus; a tiny undulated area can be seen in B. diegoi only, where the femoral spine would be, if it existed.This study verified that B. azuai and B. breuili are the same species.As both names were published in the same study by the author (Bolívar, 1921) it was first thought that it should come before azuai which appears on page 524 before breuili named on page 525; however, breuili was chosen as it is better characterized, also, the genital structure of the type series is in better condition and has therefore been studied; on the other hand the genitalia of the specimens of the type series of azuai is not preserved so must have been misplaced.MALE GENITALIA.Median lobe of aedeagus (Fig. 60) folded approximately in the middle in an obtuse angle; basal half strongly robust and dorsal face sinuous, especially in basal part.Tip of style (Fig. 61) has four long sinuous setae of similar length.Internal sac of aedeagus (Fig. 59) divided into three parts: in basal region Y-shaped piece formed by two very fine symmetrical rods with a stout bean-shaped nodule articulated by a tubule with another nodule in the same shape at the tip; there are some phanerae next to the Y-shaped piece; in median region with ventral and dorsal phanerae and two very long exterior ones, in the upper area of the median region there is phaneroid complex formed by three strongly sclerotized nodules articulated amongst each other, basal ones rectilinear with inferior nocked margin; in apical region reinforcing bands expanded in basal part forming laminas that almost touch the axis of symmetry of the sac, where a tubule can be seen.FEMALE GENITALIA.Spermatheca (Fig. 62) discreetly curved from one end to the other and median region thin; both lobes differentiated, the basal one being larger and more elongate than the apical one; spermathecal duct 13-14 times longer than spermatheca becoming thinner as it gets nearer to the spermatheca forming a long stout nodule at the point of insertion with the spermatheca; also, it is larger near the bursa copulatrix, where it crosses a sclerotized disk; the spermathecal gland opens out near the insertion point with the spermatheca.

Bathysciola (Bathysciola) (schiodtei group) breuili
DISCUSSION.The specimens in the type series from the Cueva de Mendia Landa, in Arive (Navarre) were used to illustrate the genitalia.Bathysciola diegoi should be included with those in the group with a tubule or rod amongst the reinforcing bands of the internal sac of the aedeagus.The general structure of the internal sac of B. diegoi is very similar to that of B. obermaieri (Figs. 59 and 69), but there are differences in the shape and layout of some of its sclerotized pieces; there are also some differences in the median lobe and spermatheca.
-Bathysciola diegoi: in the phaneroid complex the upper phanera is a simple triangular lamina; the intermediate one is in the shape of a triangular tooth and also a flat lamina; the ventral and lateral phanerae are strongly sclerotized and robust (Fig. 59).The dorsal area of the median lobe is more sinuous and the basal part is less robust (Fig. 60).The spermatheca (Fig. 62) is shorter and robust, with the apical lobe proportionally larger; the spermathecal duct is inserted in the spermatheca by a stout elongate nodule.-Bathysciola obermaieri: in the phaneroid complex the upper phanera forms a triangular lamina with two expanded margins; the intermediate one is elongate and the upper margin expanded, protruding towards the axial line of the sac; the ventral and lateral phanerae are more slender and weakly sclerotized (Fig. 69).The dorsal area of the median lobe in ventral view is weakly sinuous and the basal part more robust (Fig. 70).The spermatheca (Fig. 72) is longer and thin, with a proportionally small apical lobe; the spermathecal duct is thin to the point of insertion with the spermatheca though there is a small expanded area just before the point of contact.MALE GENITALIA.Median lobe of aedeagus (Fig. 64) folded approximately in half, in an obtuse angle; basal half robust and dorsal face sinuous throughout.Tip of lateral style (Fig. 65) with four similar sinuous setae which are not very long.In the internal sac of the aedeagus (Fig. 63) there are some fine, strongly sclerotized phanerae next to the Y-shaped piece; in the median region the ventral phanerae are strongly sclerotized, as are the exterior ones, which are also very long; the dorsal phanerae are hyaline; the basal nodules in the phaneroid complex are slightly arquate, almost rectilinear.
DISCUSSION.The male genitalia of the single specimen from Venoix in Calvados (France) was illustrated.The number of specimens studied is not indicated in the original description.The genitalia were prepared between two glass slides and one of the lateral style was separated; it was prepared again on an acetate paper slide and placed in Canada Balsam inserted on the same pin as the specimen it belongs to.Bathysciola fauveli was descri- bed in three lines of the identification key for the species belonging to genus Bathysciola proposed by Jeannel (1924a); in this paper the mentioned author relates the new taxon to B. rugosa due to characters of its external morphology.However, B. fauveli is closely related to B. schiodtei.The internal sac of the aedeagus was studied by transparency through the tegument of the median lobe; the reason for this was that extracting the sac meant destroying the median lobe and as this is a unique specimen it was decided that all the structures should be kept in tact.When there are more specimens the necessary preparations can be made to obtain more precise information.As far as can be seen the phaneroid complex of the upper part of the median region of the internal sac appears to be very similar to that of B. schiodtei without the upper nodules present in the latter species; in any case the internal sac of B. fauveli is certainly different to that of B. rugosa.Jeannel (1924a) states that there are three setae inserted in the tip of the lateral style.This is true of the left lateral style of the aedeagus in this specimen, but the right one has four; the left lateral style may have been mutilated; so, B. fauveli is considered to have four setae at the tip of the lateral style like the other species in the genus.MALE GENITALIA.Median lobe of aedeagus (Fig. 67) folded in obtuse angle approximately in the middle; basal half strongly robust and dorsal face strongly sinuous throughout; the tip is strongly curved in the ventral part; the tip of the lateral style (Fig. 68) has two long setae and two shorter ones, all of them twisted.The internal sac of the aedeagus (Fig. 66) has a number of long, fine well sclerotized exterior phanerae; the ventral phanerae are very evident but the dorsal ones are practically invisible; the lower nodules in the phaneroid complex are fol-ded in an acute angle; amongst the reinforcing bands there are two laminas that do not come into contact, with a short tubule between them.DISCUSSION.Specimens from Arudy, Basses Pyrénées (France) were used for the illustrations.It was described as a species (Fairmaire, 1856), but added to the subspecies of B. schiodtei (Jeannel, 1910).The peculiar features observed in the internal sac of its aedeagus in this study are sufficient to restore it to its original status of species.It should be included with those species exhibiting a tubule amongst the reinforcing bands of the internal sac.The most similar of these species is B. bigerrica (B.parallela would be slightly more distant) which has the same shaped basal nodules in the phaneroid complex and an almost identical tubule amongst the reinforcing bands; in B. bigerrica there is also a rod, which does not occur in B. grandis.(Español, 1966); S-HU, Villanúa, Cueva del Rebeco, 20.8.1965, Auroux & OE leg., 1 male (Español, 1966); S-HU, Villanúa, Cueva del Rebeco, 1962-63, Español, Senent, Auroux, Dumont, Barbier & OE leg., (Escolà, 1974); S-SS, Orio, Guardetxe-Aurre'ko Leizea 1, Galán leg.(Español, 1970: subsp. rugosa;Escolà, 1974).The determination of the latter mentioned material need to be revised as its distribution area is at a considerable distance from the normal distribution of the species.MALE GENITALIA.Median lobe of aedeagus (Fig. 70) folded in obtuse angle approximately in the middle; basal half strongly robust and dorsal face sinuous throughout; tip of lateral style (Fig. 71) with four long twisted setae of similar length, sometimes very long and twisted in a complicated manner.Internal sac of aedeagus (Fig. 69) with long robust exterior phanerae, though weakly sclerotized; phaneroid complex with rectilinear basal phanerae, upper end thin and basal thick, reforcing bands widened in basal region forming plates almost touching the axis of symmetry of the sac, where a tubule can be seen; near this a stout weakly sclerotised rod stretches from the apical tip of the reinforcing bands with a pubescent distal tip.FEMALE GENITALIA.The spermathecal complex coincides with the description given for the genus, the spermatheca (Fig. 72) being long and thin.
DISCUSSION.The genitalia of both sexes was illustrated using the specimens of the type series from the Cueva de Santa Elena, Biescas (Huesca).This taxon was described as a species (Bolívar, 1918) but added to the subspecific table of B. rugosa (Jeannel, 1924a).The peculiar features observed in the internal sac of the aedeagus are sufficient to restore it to its original status of species.Like B. diegoi -the most similar species-it belongs to the subgroup with a sclerotized structure amongst the reinforcing bands of the internal sac of the aedeagus; B. rugosa is not amongst them.The comparative examination of B. diegoi in the discussion section can be consulted.
Coleopterorum catalogus pars 60: 10 Bathysciola (Bathysciola) parallela parallela (Jeannel): Perreau, 2000.Mém. de la SEF, 4: 237 MATERIAL STUDIED.The original description indicates that only two males and two females have been studied (Jeannel, 1907); a male specimen belonging to the type series was found, stored in the MNHNP general collection and labelled: 1. MALE GENITALIA.Median lobe of the aedeagus in lateral view (Fig. 74) long, weakly arcuate, uniformly decreasing from base to tip, both faces strongly sinuous; lateral style almost as long as median lobe, pointed and bearing four strongly folded setae of similar length (Fig. 75); internal sac of aedeagus (Fig. 73) with weakly sclerotized phanerae next to Y-shaped piece; ventral phanerae in median region enormous, strongly sclerotized and forming the interior margin with translucent laminas; dorsal ones tiny and also strongly sclerotized; short phanerae along sides of sac, with only the lower pointed end sclerotized; basal nodules of phaneroid complex folded in very acute angle; inte-rior margin of reinforcement bands noticeably expanded and sclerotized, prolonged, forming enormous peduncles; robust rod amongst reinforcement bands.DISCUSSION.A series of specimens from the Grotte de l'Oueil du Neéz in Rébenacq, Basses Pyrénées (France) was used to illustrate the male genitalia.The description is very superficial as it is limited to a few lines in a dichotomic key (Jeannel, 1907).Its affinities must be looked for in the species of its group which have a sclerotized structure amongst the reinforcement bands of the internal sac of the aedeagus.B. bigerrica is the most similar and there is no doubt about its close relationship, they are the only species that have in common the unusual character of a pedunculate inferior end of the reinforcing bands.Many genital characters enable its identification (Figs.51 and 73) though the difference in robustness, size and degree of sclerotization in the phanerae in the median region of the internal sac is probably the most apparent.lengths.Internal sac of aedeagus (Fig. 76) with long fine strongly sclerotized phanerae with a laminar expanded area in upper extreme; phaneroid complex with basal phanerae folded in right angle; reinforcing bands expanding in basal region forming laminas that leave an empty space in the axial region of the sac.
FEMALE GENITALIA.Spermathecal complex (Fig. 79) coinciding with the description given for the genus, the spermatheca being proportionally short and robust.
DISCUSSION.The genitalia of both sexes were illustrated based on a series of specimens from the Cueva de Orobe, Alsásua (Navarra).This taxon was described as species (Sharp, 1872), but Coiffait (1959) tended to consider it as a large size subspecies of B. schiodtei, whose females are not distinguishable from those belonging to B. grandis -at that time considered as a subspecies of B. schiodtei-.The B. rugosa males are the largest of all the group and have such depressed elytra that they are almost flat.There are no differentiated structures in the reinforcement bands of the internal sac of the aedeagus, and the appearance of the basal nodules of the phaneroid complex place this taxon close to B. breuili, the most similar species.The characters differentiating these two species are detailed in the discussion section referring to B. breuili.
MATERIAL STUDIED AND BIBLIOGRAPHICAL DATA.The original description does not indicate the number of specimens studied and the specimen or the specimens of the type series have not been found in the investigations carried out in different museums.So, apart from the interpretation of the original description (Kiesenwetter, 1850), the characterisation of the species provided by different authors who state that they have identified the species (Bolívar, 1919;Jeannel 1911Jeannel & 1924a;;Español, 1956;Coiffait, 1959) has been accepted.Numerous specimens of French origin determined by R. Jeannel and H. Coiffat deposited in the M NHNP collection, have been located and studied.The original description indicates the type locality: "Bagnières de Luchon" (Kisenwetter, 1850); different samplings carried out in the locality did not provide any specimens though some were found in a nearby locality; alter comparing them with the MNHNP specimens and verifying that it was the same species, they were used for the descriptions and illustrations in this study.The following material was studied and Iberian bibliographical references found: S-L, Bóssost (Español, 1956); S-L, Lés, III.1916, Hilaire leg.(Bolívar, 1919;Español, 1956); XI.1916, Hilaire leg.(Bolívar, 1919;Español, 1956); S-L, Vielha-Mijaran, Plan Batalher, 10.V.1999, JF leg., 3 males, stored in col.JF.; S-L, Val d' Aran, VIII.1916, Zariquiey leg. (Bolívar, 1919).The following specimens from the MNHNP general collection were studied: S-L, Baix Aran, Bossost, 16.I.1916MALE GENITALIA.The genital structures of B. schiodtei were used in the description of the genus; internal sac of the aedeagus (Fig. 80), aedeagus in lateral view (Fig. 81) and tip of the lateral style (Fig. 82).
DISCUSSION.Bathysciola schiodtei belongs to the subgroup of species lacking tubules and rod amongst the reinforcing bands of the internal sac of the aedeagus.The most similar one is B. fauveli which also has straight, nocked basal nodules in the phaneroid complex, however, B. fauveli does not have the robust elongate upper nodules present in B. schiodtei.Coiffait, 1959Bathysciola aranensis Coiffait, 1959. Ann. Spéléol., 14(1-2): 162 Bathysciola (Bathysciola)  No male from B. aranensis Coiffait, was studied; the related specimen in the description is a female and therefore it was not possible to determine the characters of its aedeagus; in the opinion of the person who described it this taxon is related to B. meridionalis (J.du Val) which also belongs to section VI of the genus Bathysciola, Jeannel (1924a).The number of specimens in the type series is not indicated in the original description: "Type: Val d'Aran,...", "Mâle inconnu."(Coiffait, 1959).The only female specimen was found in the Coiffait collection preserved at the MNHNP and is identified as a holotype labelled: 1. (hw Coiffait): Val d'Aran / Artiga de Lin / 9.54 H. Coiffait; schiodtei.80) internal sac of the aedeagus in dorsal view.81) aedeagus in lateral view.82) apical region of the lateral style.Scale bar: a-b, 0.5 mm: 80, 81.ARB, apical reinforcement band; DPM, dorsal phanerae of the medial region; LPM, lateral phanerae of the medial region; NPYP, next phanerae of the Yshaped piece; PC, phanerae complex; SN, sclerotized nodule; VPM, ventral phanerae of the medial region; YP, Y-shaped piece.
8. Male metafemurs with denticle in posterior margin.In the internal sac of the aedeagus (Fig. 54) the two upper phanerae in the phaneroide complex are superposed, one on top of the other, so that only the apical vertex of the upper one can be seen; outer phanerae in the median region thin and weakly sclerotized .
B. bigerrica, B. diegoi, B. grandis, B. parallela and B. obermaieri.In any case B. bigerrica and B. parallela are closely related as they are the only species with the unusual character of a pendunculate lower extreme of the reinforcing bands in common.Identification is made possible by numerous genital characters (Figs.51 and 73) though the robustness, size and degree of sclerotization of the phanerae of the median region of the internal sac is possibly the most apparent.
, RJ leg., 2 males and 2 females.COMPARATIVE STUDY.The male genitalia is identical to B. o. ovata.B. o. gabasensis is characterised by its high mesosternal carina which form a right or somewhat acute angle; whereas the carina in B. o. ovata is lower and clearly obtuse.