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A New Cryptic Species of Aphaenops (Coleoptera: Carabidae: Trechinae)

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A New Cryptic Species of Aphaenops (Coleoptera: Carabidae: Trechinae) Ann. soc. entomol. Fr. (n.s.), 2007, 43 (3) : 363-370 ARTICLE A new cryptic species of Aphaenops (Coleoptera: Carabidae: Trechinae) 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 beetle 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 beetles, troglobitic, allopatric speciation. ave animals 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 insects & 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 Taxonomy 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 insect (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
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