Baronniesia Fresneda, Bourdeau & Faille

Fresneda, Javier, Bourdeau, Charles & Faille, Arnaud, 2009, Baronniesia delioti gen. n. sp. n., a new subterranean Leptodirini from the French Pyrenees (Coleoptera: Leiodidae: Cholevinae), Zootaxa 1993, pp. 1-16 : 2-8

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 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.

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 ).

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.

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, 1950 b, 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.

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).

In all other Pyrenean Leptodirini the number of spines in the apex of the lateral stylus of the aedeagus varies from two to five, but they all have a penicillum, which could be inserted in a depression or not, or be inserted on a membranous lamina.

There are two main types of lateral stylus of the aedeagus among the rest of Pyrenean Leptodirini:

1) The Troglophyes type. The apex of the lateral stylus of the aedeagus with two or (more frequently) three long spines, with a dense penicillum formed by numerous short setae. This penicillum may be inserted in a cavity or directly on the integument of the apex of the stylus ( Figs. 10–12 ). In the speluncarum group of Speonomus Jeannel, 1908 the third spine is often concealed by the penicillum. The following groups have the lateral stylus of the Troglophyes type: Bathysciella Jeannel, 1906 ; Bellesia Fresneda & Hernando, 1994 ; Ceretophyes Fresneda, 1998 ; speluncarum ( Jeannel, 1947) and ehlersi ( Perreau, 2000) groups, and the subgenus Machaeroscelis Jeannel, 1924 of Speonomus ; Phacomorphus Jeannel, 1908 , including the subgenus Phacomorphoides Dupré, 1989 ( Fig. 12 ); Parvospeonomus Bellés & Escolà, 1977 ( Fig. 11 ); Perriniella Jeannel, 1910 and Troglophyes Abeille de Perrin, 1894 ( Fig. 10 ). Troglophyes is a para- or polyphyletic genus ( Fresneda et al. 2007). All the easternmost species of Troglophyes , T. bedeli Jeannel, 1906 , T. g. gavoyi Abeille de Perrin, 1894, T. g. alluaudi Jeannel, 1911, T. ludovici Chobaut, 1903 and T. oblongulus Reitter, 1908 , are closely related to Ceretophyes Fresneda, 1998 and Perriniella Jeannel, 1910 , unlike the Antrocharis group of genera and T. a. aubryi and T. a. vallierensis of the central Pyrenees.

2) The Trocharanis type. The apex of the lateral stylus of the aedeagus is with three long spines (sometimes five). One can be longer than the other two ( Fig. 13 : Speonomus curvipes subcurvipes (Abeille de Perrin, 1878)), or two longer than the third one, all three of different size ( Fig. 14 : Paraspeonomus vandeli Coiffait, 1952 ) or all subequal. A dense penicillum formed by numerous long setae inserted on the side or on a more or less developed membranous lamina is always present; in some species the long setae of the penicillum look like a pedunculus or thin extension of the membranous lamina (e.g. in S. carrerei Fourès, 1954 ). This group is formed by the genus Paraspeonomus Coiffait, 1952 , Speonomus of pyreneus and zophosinus groups ( Jeannel 1947), subgenus Metaspeonomus Coiffait, 1959 and Trocharanis Reitter, 1885 . All the genera of the southern side of central Pyrenees and coastal ridges of Catalonia ( Spain) belong to this group: Lagariella Fresneda, 2000 , Naspunius Fresneda, Hernando & Lagar, 1994 , Pallaresiella Fresneda, 1998 , Salgadoia Fresneda, 1998 , Speonomites Jeannel, 1910 , Stygiophyes Fresneda, 1998 , Trapezodirus Jeannel, 1924 and Troglocharinus Reitter, 1908 (except the three taxa of the subgenus Antrocharidius Jeannel, 1910 with a secondary loss of penicillum but with the membranous lamina).

Dilated antennae in male of Baronniesia is a character shared with Josettekia angelinae Bellés & Déliot, 1983 and Paraspeonomus vandeli Coiffait, 1952 . Josettekia angelinae has the 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 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.

Figures

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).

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

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

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.

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 ”.

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