Ann. soc. entomol. Fr. (n.s.), 2007, 43 (3) : 363-370 ARTICLE

A new cryptic species of Aphaenops (Coleoptera: Carabidae: ) from a French Pyrenean cave: Congruence between morphometrical and geographical data confi rm species isolation Arnaud Faille (1), Philippe Déliot (2) & Eric Quéinnec (3) (1) 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 Buff on, F-75005 Paris, France (2) Laboratoire souterrain du CNRS, F-09200 Moulis, France (3) Unité «Evolution & Développement», UMR 7138 «Systématique, Adaptation, Evolution», Université P. & M. Curie, 9 quai St–Bernard, F-75005 Paris, France

Abstract. A new species of cave-dwelling from the pyrenean massif, Aphaenops (Cerbaphaenops) jauzioni n. sp. (Coleoptera: Carabidae), is described. Geometric morphometric analyses based on fi ve morphological structures and comparisons with two populations of the closely related species Aphaenops (Cerbaphaenops) cerberus Dieck 1869 support the uniqueness of the new species. The specifi c status of the new taxon is also confi rmed by the male genitalia structures. A putative scenario of population isolation is proposed. Résumé. Une nouvelle espèce cryptique d’Aphaenops (Coleoptera : Carabidae : Trechinae) d’une grotte des Pyrénées françaises : la congruence entre les données morphométriques et biogéographiques confi rme l’isolement spécifi que. Aphaenops (Cerbaphaenops) jauzioni n. sp. (Coleoptera : Carabidae), une nouvelle espèce de carabique troglobie pyrénéen, est décrit. Des analyses de morphométrie géométrique réalisées sur cinq structures morphologiques et la comparaison avec deux populations indépendantes de l’espèce proche Aphaenops (Cerbaphaenops) cerberus Dieck 1869 témoignent de la différenciation de la nouvelle espèce. Le statut spécifi que du nouveau taxon est également confi rmé par la structure de l’édéage. Un scénario est proposé pour expliquer l’isolement de la population. Keywords: Aphaenops, Trechinae, cave , troglobitic, allopatric speciation.

ave are an ideal model for studies of are known from a single cave, others display a highly Cspeciation processes because of their stable pattern fragmented distribution area, correlated with their of distribution and the partial or complete reduction specifi c ecological requirement and with the structure of of gene fl ow between populations (Barr & Holsinger karstic environments. Some species are known from the 1985). Species in the genus Aphaenops Bonvouloir 1861 “Milieu Souterrain Superfi ciel” (e.g. ‘M.S.S.’; Juberthie (Coleoptera: Carabidae) are the most signifi cant & Bouillon 1983), i.e. cracks and fi ssures in the upper of the Pyrenean cave fauna. All 38 known species of part of the rock and of interconnecting spaces in scree this genus are endemic to the Pyrenean region (Coiff ait and talus. On the north-eastern part of their range they 1962). Th e adjective “aphaenopsian” is currently used are represented by the subgenus Cerbaphaenops Coiff ait, to describe characteristics of troglobitic invertebrates 1962. In order to broaden our knowledge of the genus (i.e. troglomorphic characters; Christiansen 1992). in general, and of Cerbaphaenops in particular, we Th erefore, the suffi x -aphaenops is generally used to examined new material from a number of visits made indicate the convergent morphologies of trechines well- by the authors to caves of the Haute-Garonne between adapted to caves around the world (e.g. Neaphaenops October 2003 and November 2005. Jeannel from North America, Laosaphaenops Deuve Aphaenops cerberus Dieck 1869, the most common from Laos, Mexaphaenops Bolivar & Pieltain from species of the Cerbaphaenops complex, is distributed Mexico). Aphaenops species are diversifi ed in the major along the French Pyrenean foothills and colonizes many part of the Pyrenees range. Nevertheless, distribution caves of Ariège and Haute-Garonne departments in an pattern of the genus is heterogeneous: many species area where the karstic cover is highly fragmented. In order to identify the detailed distribution pattern of the widespread species A. cerberus, we explored peripheral E-mail: [email protected], [email protected] areas where this species was recorded (Jauzion pers. Accepté le 19 avril 2007 com.)

363 A. Faille, P. Déliot & E. Quéinnec

Material and Methods Table 1. Number of specimens studied for each structures Th e morphological variability of fi ve structures have been Species Head Pronotum Elytra Antenna Edeagus studied for population of the new species from Artigouli cave A. jauzioni n.sp. 30 30 30 30 15 (Chein Dessus, Haute-Garonne, France) and two populations A. cerberus, Marbrière 31 31 31 31 28 of A. cerberus. Although numerous populations of this latter A. cerberus, Sendé 10 8 8 8 8 species were studied in the scope of a taxonomical revision of the group, the two stations chosen in this paper are Le Sendé cave (Moulis, Ariège, France) which is the type locality of A. cerberus stricto sensu and La Marbrière cave, (Montastruc-de- Salies, Haute-Garonne, France), which is the closest locality of A preliminary analysis of these specimens and of Artigouli massif. material from other collections has revealed three Th e fi ve morphological structures we measured are the next: morphospecies, one of which diff ers considerably from head, pronotum, elytra, edeagus and antenna. Th e numbers populations of Aphaenops cerberus previously described. of specimens studied for each population are indicated in the Here, we present the description of a new species tab. 1. belonging to the subgenus Cerbaphaenops and discuss All images were taken using a Kappa DX30 digital camera the characters that distinguish it from A. cerberus using mounted on a Leica MZ16 stereomicroscope. All measurements datasets of multiple morphological landmarks. and coordinates were taken using TpsDig1 (Rohlf 2004).

Figure 1 Plots of specimens of the two populations of Aphaenops cerberus and A. jauzioni n. sp. onto the fi rst CVA plane for a, the head, b, the pronotum, c, the elytra and, d, the antenna. Aj: Aphaenops jauzioni n. sp., AcM: Aphaenops cerberus (La Marbrière), AcS: Aphaenops cerberus (Le Sendé).

364 New species of Aphaenops from Pyrenean caves

Morphometric analyses Th e three populations display signifi cant interpopulational diff erences. Moreover, the Artigouli For the head, thirteen homologous landmarks were defi ned for Cerbaphaenops species. Th ese landmarks have been used cave population exhibits high divergence in the form for a general overall analysis of cave-inhabiting trechines from of the median lobe of the male genital organ (fi g. 2). the Pyrenees (Faille 2006). For the pronotum, the elytra and Observations of this structure allow us to distinguish the genitalia we used Elliptical Fourier Analysis (EFA) that easily the Artigouli cave population from all others. It allow a study of complex structures and are particularly useful to investigate structures without well defi ned homologous is interesting to notice that the two populations of A. landmarks (Rohlf 1996; McLellan & Endler 1998). Both cerberus studied, located in the western side of the species Procrustes superimposition and EFA involve the estimation of an distribution area, show high level of diff erentiation isometric size measure (centroïd size for Procrustes approaches, of the edeagus also (fi g. 5). Based on this and on the square root of the surface bounded by the outlines). other morphological characters studied, we proposed to Concurrently, we measured traditional distances that were diff erentiate the Artigouli population in a new taxon used in the taxonomical descriptions and in a multivariate morphometric analysis of the antennal segment proportions. named Aphaenops (Cerbaphaenops) jauzioni n. sp. Th ese eleven measures are: A1–A11: length of antennal segments, AL = maximum length of aedeagus (measured from the insertion of paramers to apex); AnL = maximum length of antennae; EL = maximum length of elytra; EW = maximum width of elytra; HL = maximum length of head (measured Aphaenops (Cerbaphaenops) jauzioni Faille, from the base of the neck to apex of labrum); HW = maximum Déliot & Queinnec n. sp. width of head; HWn = maximum width of neck; M = median (Figs. 3, 4) of each measure; PA = approximate width of pronotal apex; PB = approximate width of pronotal base; PL = length of the Type material. Holotype. ♂ Artigouli cave (named also Larrieu pronotum measured along the midline; PW = maximum width cave) (43°01’03”N 0°51’34”E), Chein-Dessus, Haute-Garonne, of the pronotum; RA = antennal ratio = maximum length of France, 21.IX.2004, Déliot & Faille col. Deposited in National habitus versus maximum length of antennae. Only the antennal Museum of Natural History collection (NMNH), Paris, France. lengths A1 to A11 were analysed using log shape ratios, a Paratypes. Th irty specimens (10 ♂♂, 20 ♀♀), same collecting multivariate morphometric approach that allows a calculation data as holotype (NMNH). Additional material studied. Our of individual size measure and shape (size-free) variables colleague speleologist Georges Jauzion has collected this (Mosimann & James 1979). in two other localities of the massif near the typical locality: Puits Jozian (43°01’26’’N 0°52’35’’E), Montastruc de Salies, Statistical analyses Haute-Garonne, France, 20.X.1990: 1♂ (G. Jauzion coll.). Puits du Clot de la Bouche (43°01’23’’N 0°52’05’’E), Estadens, We used Principal Component Analysis (PCA) together with Haute-Garonne, France, 04.XII.1994: 1♂, 1♀ (G. Jauzion Canonical Variate Analysis (CVA) and Discriminant Functions coll.). Material examined for comparison. Aphaenops cerberus (DF). CVA and DF classifi cation results were calculated using ssp. cerberus Dieck 1869. 8 specimens (collection MNHN and a “leave-one-out” cross validation procedure. Canonical Variate C. Bourdeau) from Le Sendé cave (42°57’30’’N 1°07’13’’E) Analysis (CVA) are presented using the fi rst two canonical axis. Th e goal of those morphometrical studies is to investigate both the amount and the type of diff erences between populations. Since all analyses done with or without size provided similar results, those presented in this article deal only with form (i.e. size+shape). Besides PCA results did not provide any additional information and are not shown here. Morphometric and statistical analyses were carried out using Matlab v. R14 with special functions (see Baylac & Friess 2005).

Results Results provided by the morphometrical analyses are unambiguous. As shown in fi g.1a–d by the average outlines of the head, pronotum, and elytra together with the log shape ratio of the antennae, the diff erences between the three populations studied are signifi cant. “Leave-one-out” results demonstrate clearly the diff erentiation of the Artigouli population with 100% of correctly classifi ed specimens for the edeagus, the Figure 2 head and the antennae and 96.7% for the elytras Plots of specimens of the two populations of Aphaenops cerberus and A. (one single misclassifi ed specimen) and 83.3% for the jauzioni n. sp. onto the fi rst CVA plane for the edeagus antennas (fi ve misclassifi ed specimens).

365 A. Faille, P. Déliot & E. Quéinnec

Table 2. Antennae measurements of Aphaenops jauzioni male holotype. Th e ratio is length of the fi rst antennomere versus length of each antennomere Antennomeres A1 A2 A3 A4 A5 A6 A7 A8 A9 A10 A11 Lenght (mm) 0.23 0.30 0.40 0.42 0.43 0.41 0.38 0.32 0.29 0.27 0.37 Ratio A/A1 1.0 1.3 1.7 1.8 1.9 1.8 1.7 1.4 1.3 1.2 1.6

(= “grotte d’Aubert”, Ariège, France, collected in April 2003 by C. Bourdeau, P. Déliot and A. Faille. Aphaenops cerberus ssp. bruneti Jeannel 1926 from La Marbrière cave (43°00’35’’N 0°54’38’’E), Montastruc-de-Salies, Haute-Garonne, France. Diagnosis. Th is species is readily distinguished from the closest species A. cerberus by its aedeagus moderately arcuated and gradually acuminate. Description. Length: 4.27–4.47 mm; M 4.4 mm (from apical margin of clypeus to apices of elytra). Head. Long (HL 0.93–1.00 mm; M 0.96 mm) and narrow, widest at the anterior two-fi fths, and gradually narrowing just behind the neck constriction shallow but distinct and cylindrical; HL/HW 1.3–1.5 (M 1.4), HL/PL 1.22–1.25 (M 1.23); neck convex, ring-shaped, with the sides either straight or slightly curved and with the anterior constriction about three-sevenths as wide as the widest part of the head; dorsum convex, with gently convex frons and rather depressed supraorbital area; frontal furrows unevenly impressed, long and somewhat deep, slightly bisinuate before middle at the level of the anterior pair of supraorbital pores, gradually divergent in the posterior part and obsolete before the level of posterior supraorbital pores; posterior pair of supraorbital pores located far from cervical furrows, just behind the neck constriction; microsculpture fi ne but distinct, consisting of mostly wide, partially isodiametric reticulation; genae gently convex, a little more so at the posterior parts, sparsely covered with short pubescence; labrum either nearly straight or slightly emarginate at the apex, either straight or slightly bisinuate at the median part, sexsetosae as usual; mandibles falciform moderately arcuate inwards at the acute apex, right mandible in the so-called ‘bidentate’ condition, with fairly wide retinaculum and left one with only one distinct teeth. Clypeus small, quadrisetosae. Mentum totally fused with submentum, labial suture not perceptible throughout, with a small protrudent tooth in apical emargination; submentum with a transverse row of eight setae. Antennae long (AnL = 3.81 mm) and slender though not exceeding elytral apex even in male (RA = 1.15), reaching apical tenth of elytra in male, apical seventh to eight of elytra in female; scape short, about fi ve-sevenths as long as pedicel, which is only two-fi fths as long as segment 3, segments 5–10 gradually decreasing in length towards apex, all cylindrical and about nine times as long as wide in segment 5, segment 10 about as long as 2, terminal segment about as long as segment 7 and about a fi fth as wide as scape (see Tab. 2). Th orax. Elongated (PL 0.69–0.75 mm, M = 0.71 mm), subovate, a little shorter but evidently narrower than head, widest at about two-fi fths from base, and more gradually narrowing towards apex than towards base; PW/HW 0.8–1.1 (M 0.9), PL/PW 1.1–1.3 (M 1.26); apical third of propleura visible from above. Pronotum elongate, widest at about middle and narrowing more towards the base than towards apex; PL/ Figure 3 PB 1.7–1.9 (M 1.8), PL/PA 1.55–1.7 (M 1.68); sides fi nely Aphaenops (Cerbaphaenops) jauzioni n. sp., male habitus from the Artigouli bordered throughout, very feebly arcuate at the middle and very cave (Haute–Garonne). briefl y but visibly sinuate just before hind angles; median line fi ne though distinct; front angles rounded or obtuse though

366 New species of Aphaenops from Pyrenean caves distinct, but hind angles usually subrectangular though not so lobe; basal part gently curved in ventral side, with large basal sharply defi ned, sometimes narrowly rounded at the corners; orifi ce whith sides widely emarginated; sagittal aileron very two pair of marginal setae present, the anterior pair situated small and hyaline but present; viewed dorsally, apical lobe just before the widest part, at about apical two-sevenths and the nearly symmetrical, gradually narrowing towards the tip, which posterior pair at the two-sevenths from the base; apex nearly is obtusely subangulate; viewed laterally apical lobe slender, straight or slightly arcuate; dorsum convex, steeply declivous at strongly refl exed at the apical portion, and abruptly blunt at the the antero-laterally; microsculpture sharply impressed, mostly extremity. Inner sac armed with a subspatulate copulatory piece consisting of transverse meshes, partially of transverse lines; just inside thin and hyaline apical orifi ce, about two-ninths as apical transverse impression. Abdomen. Elytra oblong-oval long as the aedeagus. Paramers fairly large with elongate apical (EL 2.66–2.75 mm, M 2.7 mm), imperfectly fused together, parts but devoid of ventral apophysis even on the left one, each more than twice as wide as prothorax (EW/PW 2.2–2.4, usually bearing two apical setae. M 2.3), nearly twice as long as wide (EL/EW 0.45–0.6, M 0.55), widest at about the middle and contracted at the bases; Variability. Specimens of A. jauzioni n. sp. show few variations shoulders eff aced, with long, very oblique prehumeral borders; in setal position. Only two specimens of twelve observed for sides narrowly bordered throughout, unevently ciliated, not this character display a slightly antero-posterior displacement of serrulate; dorsum strongly convex, steeply declivous at the sides the second and/or third macrochaetae of the discal group. Such and apical part, more gradually declivous towards bases and displacements, or even losses, are frequent in some species of distinctly depressed on each side of suture, with the area around Cerbaphaenops (Piochard de la Brûlerie 1872; Jeannel 1928). As scutellum moderately convex; surface irregularly subvariolate, variability is low in this population it indicates the species has a subopaque and deprived of pubescence; striation eff aced more homogenous morphology compared to some populations altogether, though vague longitudinal depressions probably of the closely related species A. (C.) cerberus (e.g. in form of the representing the vestiges of inner striae sometimes perceptible head, form of the median lobe of the aedeagus, number of styles on the disc; three setiferous dorsal pores (macrochaetes) present of paramers). on the site of stria 3, preapical pore present on the site of stria Etymology. Th is species is dedicated to Georges Jauzion, 3 located on the apical declivity near to apex, but more distant indefatigable explorer of the eastern pyrenean caves, with all from apex than from suture. Marginal umbilicate pores not our thanks for his valuable collaboration. aggregated; pore 1 of the humeral set widely removed inwards Distribution. Th is new species occurs in the small hilly area and backwards, pores 2 to 4 ranged equidistantly, pore 2 at the north-western side of the Estelas-Arbas massif between adjoining marginal gutter but the third and fourth pores are 600 and 800 meters in altitude. In a straightline, Larrieu cave isolated and moderately distant from the gutter; middle set of marginal umbilicate pores (5 and 6) displaced inwards, the sixth is only about 2.6 km from Marbrière cave (Arbas, Haute- pore lying at about apical fourth and a little more distant from Garonne) which represents the western limit of the distribution marginal gutter than pore 5; apical set of marginal umbilicate of Aphaenops cerberus. pores also displaced posteriorly, the eighth pore being located Artigouli cave is a fairly dry cavity restructured by mining at apical thirteenth to fourteenth, evidently behind the level of working (Jauzion 1985). Surprisingly, cave beetles of the genus preapical pore. Aedeagus (Fig. 4). Small (AL = 0.5 mm) though Speonomus (Leiodidae: Leptoderini) which are common in lightly sclerotized; moderately arcuated excepted for basal part, numerous cavities of the Estelas-Arbas massif, are lacking here. gently depressed, and gradually acuminate from behind middle Aphaenops jauzioni is probably present in all favourable karstic in profi le, with fairly large basal part and long fl attened apical environment of the Bois de Th ouas massif localized between

Figure 4 Male genitalia of Aphaenops (Cerbaphaenops) jauzioni n. sp., aedeagus of the holotype male from Artigouli cave (B) and, for comparison, (A) Aphaenops (Cerbaphaenops) cerberus from Sainte–Catherine cave (Balaguères, Ariège) left lateral view.

367 A. Faille, P. Déliot & E. Quéinnec

Figure 5 Visualization of the outlines (after extraction of size parameters) of the aedeagus (above), pronotum and elytra (below) for two populations of A. cerberus and A. jauzioni n. sp. A, A. jauzioni n. sp.; B, A. cerberus (La Marbrière); C, A. cerberus (Le Sendé). Th e number of specimens is given under each outlines.

Figure 6 Schematic distribution map of the Aphaenops cerberus subspecies and A. jauzioni n. sp.. In grey: karstic areas; Points: localities of Cerbaphaenops species. 1. Artigouli cave– 2.–. La Marbrière cave (Montastruc–de–Salies, Haute–Garonne)– 3. Le Sendé cave (Moulis, Ariège) – 4. Sainte–Catherine cave (Balaguères, Ariège); (10 km between two crosses).

368 New species of Aphaenops from Pyrenean caves

Estadens and Montastruc-de-Salies (Haute-Garonne), where which occured prior to the world last glacial maximum it replaces Aphaenops cerberus. Th e Bois de Th ouas represents (LGM), some 40,000 to 38,000 years ago; see Garcia- a micro-karstic area of 3.7 square kilometers consisting of Ruiz et al 2003) the temperate species, that are today Jurassic black dolomites and dolomitic limestone (mainly Kimmerdgien), which is completely separated by Pliocene widespread in western Europe, may have survived sediments from large cretaceous limestones situated along the in small and climatically favourable areas located in Estelas-Arbas massif (Fig. 6). the southern peninsulas of Iberia, Italy and Balkans. Th e case of troglobitic fauna is somewhat diff erent. Discussion Aphaenops species were located in small favourable Aphaenops jauzioni is clearly distinct from previously spots within the Pyrenean region. After progressive described Cerbaphaenops species by the peculiar climatic warming, some of the “surviving” populations morphology of male genitalia (Fig. 5). Its male genitalia expanded into endogean environments or persisted are similar to those of A. cerberus in overall form but by shifting altitude into nival environments. Th is diff ers from the latter in the shape of the median lobe, hypothesis, referred as “Climatic relict hypothesis” in particular in that the apical lobe is wider (Figs. (Peck & Finston 1993), could be proposed to explain 4–5). Aphaenops cerberus and A. jauzioni have many the isolation and diff erentiation of A. jauzioni. characters in common, i.e. form of labium, position Acknowledgments. We are grateful to Charles Bourdeau (Toulouse, France), Gabriel Corbaz (Cozzano, France), Eric of marginal setae on elytra, general shape of head Ollivier (Le Havre, France), Sébastien de Rossi (Antony, France) and pronotum and dilatation of male protarsomeres. and Christian Vanderbergh (Les Clayes ss Bois, France) for Aphaenops jauzioni diff ers from A. cerberus in the their help with collecting samples during fi eld investigations. relative length of antennae, its highly convex elytra Th anks to Michel Baylac (Paris, France), Louis Deharveng with sides moderately arcuate near shoulders and more (Paris, France) and Th ierry Deuve (Paris, France) for there useful advice, and Penelope Greenslade (Australian National elongated pronotum (Fig. 5). University, Department of Botany and Zoology) who has read Aphaenops (Cerbaphaenops) cerberus is one of the this paper. most widespread species of the genus. 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