Zootaxa, Baronniesia delioti gen. n. sp. n., a new ... - CiteSeerX

Zootaxa 1993: 1–16 (2009) www.mapress.com / zootaxa/

ISSN 1175-5326 (print edition)

Article

Copyright © 2009 · Magnolia Press

ZOOTAXA ISSN 1175-5334 (online edition)

Baronniesia delioti gen. n. sp. n., a new subterranean Leptodirini from the French Pyrenees (Coleoptera: Leiodidae: Cholevinae) JAVIER FRESNEDA1, CHARLES BOURDEAU2 & ARNAUD FAILLE3 1

Javier Fresneda, Ca de Massa, 25526 Llesp - El Pont de Suert, Lleida, Spain. E-mail: [email protected] Charles Bourdeau, 5 chemin Haut-Fournier, F-31320 Rebigue, France. E-mail: [email protected] 3 Arnaud Faille, C.P.50, UMR 5202 du CNRS / USM 601 "Origine, Structure et Evolution de la Biodiversité", Muséum National d'Histoire Naturelle, Département Systématique et Evolution, Bât. Entomologie, 45 rue Buffon, F-75005 Paris, France. E-mail: [email protected] 2

Abstract Baronniesia delioti Fresneda, Bourdeau & Faille gen. n. & sp. n., tribe Leptodirini (Coleoptera, Leiodidae, Cholevinae), is described from the French central Pyrenees (Hautes-Pyrénées, Esparros, subterranean river of Artigaléou-Arodets). The genus is remarkable for the large body size and the strong sexual dimorphism in antennae of its type species. The structure of the aedeagus and the inner sac, with two additional strongly curved feather-like structures, forming an inverted “V”, and the lateral styles with only three spines without other structures, indicates affinities between the new genus and the Antrocharis group, diversified in the eastern part of the Pyrenean range. The limestone area in the central Pyrenees where this new genus was discovered appears to be a contact area between the eastern and western faunas of the Pyrenean cave beetles, both for Leiodidae: Leptodirini and Carabidae: Trechini. Key words: Baronniesia delioti gen. n. sp. n., Leiodidae, Cholevinae, Leptodirini, subterranean environment, Pyrenees, France

Introduction The Pyrenees is known to be one of the world’s hot-spots of subterranean biodiversity (Culver et al. 2006). Particularly, two radiations of subterranean Coleoptera occur along the Pyrenean chain: Leiodidae: Cholevinae: Leptodirini and Carabidae: Trechini. Leptodirini are known in the Pyrenees and associated mountain chains by 222 taxa (species and subspecies) included in 28 genera, all of them endemic of this area. This group has fascinated biospeologists and evolutionary biologists in general since very early, due to its very strong ecological requirements, isolation of populations, morphological differentiation and multiple examples of character convergence. The Pyrenean Leptodirini have been studied in detail by many authors, among them Jeannel (1911, 1924), Dupré (1989, 1990) and Fresneda & Salgado (2006). A recent study suggested the monophyly of all Pyrenean Leptodirini (Fresneda et al. 2007). During biospeleological explorations in the massif of the Baronnies (Central Pyrenees, France), two of us (CB, AF) discovered an unexpected species of Leptodirini, which should be considered as the only representative of a new genus close to a group of genera with a more eastern distribution.

Material and methods The terminology of the structures of the inner sac of the aedeagus (Figs. 5–6) follows Jeannel (1924), Bellés (1984), Dupré (1992) and Fresneda (1998): Accepted by V. Gusarov: 22 Dec. 2008; published: 2 Feb. 2009

1

1) Basal area: FAPY “fibras apicales de la pieza en Y”, apical fibers of PY (Fresneda 1998) NL “nodule de liaison”, connection nodule (Dupré 1992) PY “piece en Y” (Jeannel 1924) 2) Median area: BDAM “bolsas dorsales del ápice de la región media”, dorsal sacs of apex of median area (Fresneda 1998) FDM “faneres dorsals de la regió mitjana”, dorsal feather-like structures (Bellés 1984) FVM “faneres ventrals de la regió mitjana”, ventral feather-like structures (Bellés 1984) IVP, inverted V phanerae or feather-like structures 3) Apical area: BRA “bandelette de renforcement apical”, reinforcement bands (Jeannel 1924)

Baronniesia Fresneda, Bourdeau & Faille gen. n. Type species: Baronniesia delioti Fresneda, Bourdeau & Faille sp. n.

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. 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

2

· Zootaxa 1993 © 2009 Magnolia Press

FRESNEDA ET AL.

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.

―

FIGURES 1–2. Habitus of Baronniesia delioti gen. n. sp. n. 1 male; 2

―female.

The Antrocharis group was characterized by the following characters (the terminology of the structures of the inner sac of the aedeagus follows Jeannel 1924, Bellés 1984, Dupré 1989, 1992 and Fresneda 1998): 1) The median lobe of the aedeagus in lateral view is curved forming right or obtuse angle, with a strong depression or dorsal sinuosity in the apical third. Until now this character was considered a synapomorphy of the group. The median lobe of aedeagus of Baronniesia in lateral view is curved forming an obtuse angle, but without any depression or sinuosity, decreasing regularly from base to apex, and strongly curved in the central part (Fig. 4). 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). BARONNIESIA FROM THE FRENCH PYRENEES

Zootaxa 1993 © 2009 Magnolia Press ·

3

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.

FIGURES 3–4. Aedeagus of Baronniesia delioti gen. n. sp. n. 3—dorsal view; 4—lateral view.

The Antrocharis group includes four genera: Antrocharis (two species), Gesciella Giachino & Guéorguiev, 1989 (monospecific genus), Paratroglophyes Fourès, 1954 (two species) and Troglophyes Abeille de Perrin, 1894 in pars sensu Fresneda et al. 2007: T. aubryi Coiffait, 1953 with the subspecies T. a. vallierensis Coiffait, 1953 (Fresneda et al. 2007). Coiffait (1953) indicates that the two subspecies of Troglophyes aubryi have the apex of the styles each with three spines and a penicillum composed of 7–8 setae. We did not see a penicillum in the numerous specimens from the Anglade cave (locus typicus of T. aubryi aubryi) and various MSS (“Milieu souterrain superficiel” Juberthie et al. 1980) in the area; all examined specimens have only three spines without penicillum (Fig. 9). Eskualdunella delespierrei Coiffait, 1950b, an enigmatic species from the French Basque country, known by a single specimen, seems to have a similar structure of the stylus apex, as the members of the Antrocharis group: three spines without penicillum (Coiffait 1950b: Fig. 11; 1955). But in Eskualdunella one of the spines is longer and more robust than the two others.

4


FRESNEDA ET AL.

FIGURES 5–6. Inner sac of aedeagus of Leptodirini. 5—Baronniesia delioti gen. n. sp. n. and 6—Antrocharis querilhaci dispar Abeille de Perrin. BDAM, dorsal sacs of apex of median area; BRA, reinforcement bands; FDM, dorsal feather-like structures; FVM, ventral feather-like structures; NL, connection nodule; FAPY, apical fibers of PY; IVP, inverted “V” feather-like structures; PY, “piece en Y”.

In the genus Bathysciola Jeannel, 1910, the species of ovata group (Perreau 2000, Fresneda & Salgado 2006) have the apex of the stylus with only three spines, but the mesoventral keel is extended over the metaventrite, and transverse striae on the elytra are missing. Those characters separate the ovata group from the Antrocharis group and Baronniesia. The species of madoni and zariquieyi groups are closely related to ovata group (Fresneda et al. 2007), and have a variable number of spines in the lateral stylus of the aedeagus (between 5 and 14). In the phylogenetic reconstruction by Fresneda et al. (2007) these groups of Bathysciola are not closely related to the Antrocharis group.The other Pyrenean Bathysciola (lapidicola, meridionalis, larcennei and schiodtei groups) have four spines in the apex of the lateral stylus of the aedeagus (the only exception is one species of the meridionalis group, B. finismillennii Fresneda & Salgado, 2006 with nine spines), and belong to a clade of Leptodirini restricted to the Spanish Basque country (see Fresneda et al. 2007): Josettekia Bellés & Déliot, 1983, Speocharidius Jeannel, 1919, Euryspeonomus Jeannel, 1919 (all with four spines) and Aranzadiella Español, 1972 (with about thirty spines). BARONNIESIA FROM THE FRENCH PYRENEES


5

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.

FIGURES 7–14. Apices of lateral styles of aedeagus of Leptodirini. 7—Baronniesia delioti gen. n. sp. n.; 8—Paratroglophyes jeanneli Coiffait; 9—Troglophyes aubryi aubryi Coiffait; 10—Troglophyes gavoyi Abeille de Perrin; 11—Parvospeonomus delarouzeei Fairmaire; 12—Phacomorphus (Phacomorphoides) sioberi sioberi (Coiffait); 13—Speonomus curvipes subcurvipes Abeille de Perrin; 14—Paraspeonomus vandeli Coiffait.

6


FRESNEDA ET AL.

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 9th (especially) and 10th 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

BARONNIESIA FROM THE FRENCH PYRENEES


7

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 4.0b10 (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.

Baronniesia delioti Fresneda, Bourdeau & Faille sp. n. (Figs. 1–5, 7, 15–16) Type locality. France, Hautes-Pyrénées, Esparros, subterranean river of Artigaléou-Arodets. Type series. Holotype: FRANCE: %, Esparros (massif of Baronnies), subterranean river of ArtigaléouArodets, 10.V.2007, Bourdeau & Faille leg. (Muséum National d’Histoire Naturelle, Paris). Paratypes: 33%% and 61&&, 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. TABLE 1. Measurements of antennal articles of the holotype male and a single female paratype (40 units = 1 mm). Article Male Female

I

II

III

IV

V

VI

VII

VIII

IX

X

XI

Length

17

10

8

10

12

11

13

12

15

7

14

Width

6

4

4

6

8

10

14

14

13

6

5

Length

13

10

8

9

10

10

11

8

11

7

12

Width

4

3

3

3

4

4

5

3

5

5

5

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.

8


FRESNEDA ET AL.

FIGURES 15–16. Baronniesia delioti gen. n. sp. n. 15—urite VIII of female; 16—female genital segment with spermathecal complex.

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. Etymology. This new species is dedicated to our entomologist colleague and friend Philippe Déliot, who died recently. 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, 1950a 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 1950a). 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.



9

FIGURE 17. Phylogram of the strict consensus of the 330 trees obtained after successive reweighting of the characters (in the tree, the character weights were reset to unity, so the scale of the branch lengths is uniform across the tree). Taxa with identical character states were lumped in a single terminal taxon except the genera of the Antrocharis group (see Appendix 2).

10


FRESNEDA ET AL.

FIGURE 18. Distribution of troglobitic Trechini in Pyrenees. Aphaenops (Aphaenops) Bonvouloir (continuous line); Aphaenops (Cerbaphaenops) Coiffait (doted line).

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). Baronniesia delioti is sympatric with Speonomus bastideus (speluncarum group) and three species of Trechini: Hydraphaenops elegans Gaudin, 1945, Cerbaphaenops aff. aeacus Saulcy, 1864 and Geotrechus discontignyi canteti Cabidoche, 1967. The subterranean river of Artigaléou-Arodets is the third known locality of Speonomus bastideus, which was so far only known from the caves of Labastide and Diable Rouge, both in Banios, Hautes-Pyrénées. It may be expected that the two main unrelated groups of cave Coleoptera (Leptodirini and Trechini) living in the same geographical area show some similarities in their patterns of distribution and colonization of the subterranean environment. Both live in the same underground ecosystem, characterized by strict and constant features, and share the area affected by the same climatic conditions and a particular paleogeographic history. 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 BARONNIESIA FROM THE FRENCH PYRENEES


11

FIGURE 19. Map of the Pyrenees with distribution areas of taxa with different models of lateral styles of the aedeagus in Leptodirini. The Antrocharis type (continuous line): Antrocharis querilhaci dispar Abeille de Perrin (a). The Troglophyes type (dots and dashes): Phacomorphus (Phacomorphus) fratyi (Dupré) (b), Speonomus (Speonomus) ere Escolà & Fresneda (c), Perriniella faurai Jeannel (d), Bellesia espanyoli (Auroux & Bellés) (e) and Parvospeonomus urgellesi (Español) (f). The Trocharanis type (doted line): Speonomus (Speonomus) curvipes subcurvipes (Abeille de Perrin) (g), Salgadoia brieti (Jeannel) (h) and Troglocharinus (Troglocharinus) ferreri ferreri (Reitter) (i).

12


FRESNEDA ET AL.

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. 1D), the two species of the ehlersi group, S. ehlersi Abeille de Perrin, 1872 and S. opisthonoxus Gers & Dupuis, 1988 (Gers & Dupuis 1988: Fig. 3C), 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.

Acknowledgements We thank Ignacio Ribera (MNCN, Madrid) for his help with phylogenetic analyses and comments on the manuscript, E. Ollivier (Le Havre), E. Queinnec (Paris) and J-P. Cassou (Lourdes) for their help during the field work. We wish to acknowledge Mark Stevens (Massey University, New Zealand) who accepted to read the manuscript. This work was partly funded by project CGL2007-61943/BOS to A. Cieslak (MNCN, CSIC).

References Abeille de Perrin, E. (1872) Études sur les Coléoptères cavernicoles suivies de la description de 27 Coléoptères nouveaux français. Notes sur les insectes cavernicoles de l’Ariège. Marius Olive, Marseille, 41pp. Abeille de Perrin, E. (1878) Notes sur les Leptoderites. Bulletin de la Société d’Histoire naturelle de Toulouse, 12, 144–155. Abeille de Perrin, E. (1894) Un genre nouveau et quatre espèces nouvelles de Coléoptères français. Annales de la Société entomologique de France, 63, 25–28. Bellés, X. (1984) Estudi de l’armadura genital en les poblacions de Speonomus (Parvospeonomus) delarouzeei s. l. i S. (P.) vilarrubiasi Zariquiey (Col., Catopidae, Bathysciinae) que conviuen a la Cova del Far (Susqueda, La Selva). Exploracions, 8, 7–13. Bellés, X. & Déliot, P. (1983) Nouveaux Bathysciinae (Coléoptères Catopidae) des Pyrénées françaises et espagnoles. Mémoires de Biospéologie, 10, 237–243. Bellés, X. & Escolà, O. (1977) Nuevos datos sobre los Speonomus del grupo delarouzeei (Fairmaire, 1860): Parvospeonomus subgen. nov. (Col. Bathysciinae). Speleon, 23, 33–37.



13

Bolívar, C. (1921) Silphidae. In: Bolívar, C. & Jeannel, R. Coleópteros cavernícolas nuevos de las provincias vascas. Boletín de la Real Sociedad española de Historia natural, tomo del Centenario, 50, 509–539. Bonvouloir, H. de (1862) Description d’un genre nouveau et de deux espèces nouvelles de coléoptères de France. Annales de la Société entomologique de France, (4)1 [1861], 567–571. Cabidoche, M. (1965) Sur les Aphaenops du groupe rhadamanthus (Col. Carab.). Annales de Spéléologie, 20(4), 523–528. Cabidoche, M. (1967) Coléoptères troglobies et endogés des Pyrénées Occidentales (Col. Carab.). Annales de Spéléologie, 22(3), 647–658. Chobaut, A. (1903) Description de deux Coléoptères cavernicoles nouveaux du Midi de la France. Bulletin de la Société entomologique de France, 8, 263–265. Coiffait, H. (1950a) Un Speonomus nouveau de la vallée d’Aure (Hautes-Pyrénées). Notes Biospéologiques, 5, 63–64. Coiffait, H. (1950b) Trois Bathysciites nouveaux de Larrau (Basses-Pyrénées). Notes Biospéologiques, 5, 65–70. Coiffait, H. (1952) Note sur les Bathysciites. Description d’un genre nouveau et de formes nouvelles. Notes Biospéologiques, 7, 41–44. Coiffait, H. (1953) Quatre nouveaux coléoptères cavernicoles du massif du Montvallier (Ariège). Notes Biospéologiques, 8, 27–32. Coiffait, H. (1955) Nouveaux coléoptères cavernicoles. Notes Biospéologiques, 9(2) [1954], 99–117. Coiffait, H. (1959) Note sur les Bathysciitae de la région pyrénéenne et de Catalogne. Annales de Spéléologie, 14(1–2), 159–179. Culver, D.C., Deharveng, L., Bedos, A., Lewis, J.J., Madden, M., Reddell, J.R., Sket, B., Trontelj, P. & White, D. (2006) The mid-latitude biodiversity ridge in terrestrial cave fauna. Ecography, 29, 120–128. Dupré, E. (1989) Bathysciinae (Col. Catopidae) des Pyrénées Occidentales: I. Étude du sac interne et morphologie comparée des Speonomus de la région de St-Jean-Pied-de-Port, redéfinition du sous-genre Phacomorphus et création du sous-genre Phacomorphoides, considérations phylogéniques et biogéographiques locales, considérations générales sur la systématique des Bathysciinae. Ikartzaleak, 13, 55–99. Dupré, E. (1990) Bathysciinae (Col. Catopidae) des Pyrénées occidentales: II. Étude et morphologie comparée du sac interne de l’édéage des Speonomus et genres voisins des Sierras de Navarre et Guipuzkoa. Considérations phylogénétiques et biogéographiques locales. Ikartzaleak, 14, 57–72. Dupré, E. (1992) Analyse comparée du sac interne des Bathysciinae (Col., Catopidae): intérêt taxonomique, considérations sur l’évolution fonctionnelle du sac. Mémoires de Biospéologie, 19, 169–186. Escolà, O. & Canció, E. (1983) Un nou Aphaenops de la Ribagorça (Coleoptera Trechinae). Speleon, 26–27, 75–77. Español, F. (1972) Nuevos datos sobre los Bathysciinae de Guipúzcoa (Col. Catopidae). Eos, 47, 59–66. Faille, A. (2006) Endémisme et adaptation à la vie cavernicole chez les Trechinae Pyrénéens (Coleoptera: Carabidae). Approches moléculaire et morphométrique. Thèse inédite. Muséum National d’Histoire Naturelle, Paris. 319 pp. Fourès, H. (1954) Diagnose préliminaire de nouveaux Coléoptères cavernicoles de la région du mont Valier (Ariège). Bulletin de la Société linnéenne de Lyon, 23, 3–5. Fresneda, J. (1998) Revisión de los géneros de Leptodirinae de la sección Speonomus del sur de Pirineos: géneros Antrocharidius Jeannel 1910, Perriniella Jeannel 1910, Speonomus Jeannel 1908, Troglocharinus Reitter 1908 y Troglophyes Abeille 1894. (Coleoptera, Cholevidae). Mémoires de Biospéologie, 25, 53–86. Fresneda, J. (2000) Lagariella nom. n. para substituir Lagaria Fresneda, 1998 homónimo más moderno de Lagaria Dallas, 1852 (Coleoptera, Cholevidae, Leptodirinae). Mémoires de Biospéologie, 27, 147–148. Fresneda, J. & Hernando, C. (1994) Descripción de Bellesia n. gen. (Col. Cholevidae) del Pirineo de Huesca (España) y consideraciones sobre las estructuras del saco interno del edeago. Mémoires de Biospéologie, 21, 57–62. Fresneda, J., Hernando, C. & Lagar, A. (1994) Contribució al coneixement de Speonomus eseranus Lagar, 1974 (Coleoptera, Cholevidae, Leptodirinae) i descripció del subgénere Naspunius nov. Bolletí de la Societat d’Història natural de les Balears, 37, 109–116. Fresneda, J. & Salgado, J.M. (2000) Revisión de los géneros de Leptodirinae de la Sección Speonomus del sur de los Pirineos. II: géneros Aranzadiella Español, 1972, Euryspeonomus Jeannel, 1919, Kobiella Español y Bellés, 1980 y Speocharidius Jeannel, 1919 (Coleoptera, Cholevidae). Mémoires de Biospéologie, 27, 41–52. Fresneda, J. & Salgado, J.M. (2006) The genus Bathysciola Jeannel, 1910 in the Iberian Peninsula and Pyrenees. Taxonomic revision of the sections IV, VI and VII (Jeannel, 1924) (Coleoptera, Cholevidae, Leptodirinae). Graellsia, 62(1), 25–74. Fresneda, J., Salgado, J.M. & Ribera, I. (2007) Phylogeny of Western Mediterranean Leptodirini, with an emphasis on genital characters (Coleoptera: Leiodidae: Cholevinae). Systematic Entomology, 32, 332–358. Gaudin, L. (1945) Troglobies nouveaux des Pyrénées françaises (Col., Carabidae). Miscellanea Entomologica, 42(1), 19–24. Gers, C. (1989) Speonomus orgibetensis, n. sp., coléoptère Bathysciinae troglobie du milieu souterrain superficiel. Annales de la Société entomologique de France (N.S.), 25(1), 105–116. Gers, C. & Dupuis, M. (1988) Une nouvelle espèce remarquable du genre Speonomus (Coleoptera, Bathysciinae).

14


FRESNEDA ET AL.

Mémoires de Biospéologie, 15, 167–173. Giachino, P.M. & Guéorguiev, V.B. (1989) Un nouveau genre de Bathysciinae des Pyrénées françaises (Coleoptera, Bathysciinae). Revue suisse de Zoologie, 96(2), 403–410. Jeannel, R. (1906) Contribution la faune cavernicole des Basses-Pyrénées (Col.). Bulletin de la Société entomologique de France, 11, 22–24. Jeannel, R. (1908) Étude sur le genre Speonomus Jeann. (Silphides cavernicoles pyrénéens) et sur sa distribution géographique. L'Abeille, 31, 57–102. Jeannel, R. (1910) Biospeologica XIV. Essai d'une nouvelle classification des Silphides cavernicoles. Archives de Zoologie expérimentale et générale, 45(1), 1–48. Jeannel, R. (1911) Biospeologica XIX. Révision des Bathysciinae (Coléoptères, Silphides). Morphologie, distribution géographique, Systématique. Archives de Zoologie expérimentale et générale, (5)7, 1–641. Jeannel, R. (1919) Bathysciinae nouveaux des Pyrénées espagnoles. Boletín de la real Sociedad española de Historia natural, 19, 129–137. Jeannel, R. (1924) Monographie des Bathysciinae (Col. Catopidae). Archives de Zoologie expérimentale et générale, 63(1), 1–436. Jeannel, R. (1928) Monographie des Trechinae. Morphologie comparée et distribution d’un groupe de Coléoptères. Troisième Livraison : les Trechini cavernicoles. L’Abeille, 35, 1–808. Jeannel, R. (1947) Coléoptères cavernicoles nouveaux avec une étude de la phylogénie des Speonomus. Notes Biospéologiques, 1(1), 83–95. Juberthie, C., Delay, B. & Bouillon, M. (1980) Extension du milieu souterrain en zone non calcaire: Description d’un nouveau milieu et de son peuplement par les coléoptères troglobies. Mémoires de Biospéologie, 7, 19–52. Perreau, M. (1989) De la phylogénie des Cholevidae et des familles apparentées (Coleoptera, Cholevidae). Archives des Sciences Genève, 39(3), 579–590. Perreau, M. (2000) Catalogue des Coléoptères Leiodidae Cholevinae et Platypsyllinae. Mémoires de la Société entomologique de France, 4, 1–460. Reitter, E. (1885) Bestimmungs-Tabellen der europäischen Coleopteren, 12 Necrophaga. Verhandlungen des naturforschenden Vereines in Brünn, [1884], 23, 3–122. Reitter, E. (1908) Dichotomische Übersicht der Blinden Silphiden-Gattungen. Wiener entomologische Zeitung, 27(2–3), 103–118. Saulcy, F. de. (1864) Faune française et européenne. Descriptions et remarques. Annales de la Société Entomologique de France, 4(4), 253–260. Swofford, D.L. (2002) PAUP*. Phylogenetic Analysis using Parsimony (* and other methods), Version 4.0b10. Sinauer Associates, Sunderland, MA.



15

Appendix 1. Data matrix used in the phylogenetic analysis. Terminal taxa with identical character states were pooled except for the genera of the Antrocharis group (see Appendix 2 for the composition of the groups, and Fresneda et al. (2007) for the definition of characters 1–34). Characters Speonomus pyreneus gr. Speonomus speluncarum gr. Paraspeonomus gr. Speocharidius gr. Euryspeonomus gr. Antrocharis Baronniesia gen. n. Gesciella Paratroglophyes Troglophyes a. aubryi/a. vallie Bathysciola series IV Speocharinus gr. Quaestus gr. Catops Speonemadus Breuilia Breuilites Espanoliella Oresigenus Q.(Amphogeus) escalerai Anillochlamys Ovobathysciola Paranillochlamys Pseudospeonomus Spelaeochlamys Patriziella Notidocharis Speonomidius Bathysciola ovata gr. T. (Antrocharidius)

0000000001111111111222222222233333333 1234567890123456789012345678901234567 11111011001000111110?0111101110111?00 11111011001000111110?0111111100111?00 11111011001000111110?0111101100111?00 11111011001000111110?0111001001111?00 11111011001000111110?0111001000111?00 11111011001000111110?0111001100111011 11111011001000111110?0111001100111011 11111011001000111110?0111001100111011 11111011001000111110?0111001100111011 11111011001000111110?0111001100111011 11111111001000111110?0111001100111100 1111111101110010001101000001000100??? 11111111011100100011010000011001001?? 00000100??0001000000?0000000?000????? 00000000??0010000000?0000000?000????? 11111111011100100000?0000001000100??? 11111111011100100010?10000011001001?? 11111111011100100000?00000011001001?? 11111111011100100011000000011001001?? 11111111011100100001010000011001001?? 1111111110110010001110000001101100??? 11111111101100100011100000011001011?? 1111111110110010001100000001101100??? 11111111001100110010?010000110010010? 11111111101000100011100000011001001?? 11111111001100100011100000011001011?? 0111101100110000001100000001000101??? 1111101100110000001100000001000101??? 11111111011000111110?0111001100111100 11111011001000111110?0111001110111?00

Appendix 2. Composition of the groups of terminal taxa sharing identical character states. Quaestus gr.: Q. (Amphogeus) cantabricus, Q. (Quaestus), Q. (Quaesticulus), Q. (Speogeus), Q. (Samanolla), Cantabrogeus, Leonesiella. Speocharinus gr: Speocharinus, Quaestus (Asturianella). Bathysciola series IV: B. madoni gr., B. zariquieyi gr. Euryspeonomus gr: Aranzadiella, Bathysciola meridionalis gr., B. lapidicola gr., B. schiodtei gr., Euryspeonomus, Josettekia. Speocharidius gr.: Speocharidius, Bathysciola larcennei gr. Paraspeonomus gr.: Bellesia, Lagariella, Paraspeonomus. Speonomus pyreneus gr.: Naspunius, Pallaresiella, Salgadoia, Speonomites, Stygiophyes, Trapezodirus, Troglocharinus (Troglocharinus), Speonomus pyreneus gr. Speonomus speluncarum gr.: Ceretophyes, Parvospeonomus, Perriniella, Phacomorphus, Speonomus (Batinoscelis), Speonomus speluncarum gr., Troglophyes partim.

16


FRESNEDA ET AL.