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New Data on Troglorites Breuili Jeannel 1919 (Coleoptera: Carabidae: Pterostichini): a Hypogean Iberian Species with Description of a New Subspecies Vicente M

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New Data on Troglorites Breuili Jeannel 1919 (Coleoptera: Carabidae: Pterostichini): a Hypogean Iberian Species with Description of a New Subspecies Vicente M Ann. soc. entomol. Fr. (n.s.), 2010, 46 (3–4) : 537-549 ARTICLE New data on Troglorites breuili Jeannel 1919 (Coleoptera: Carabidae: Pterostichini): a hypogean Iberian species with description of a new subspecies Vicente M. Ortuño (1), Javier Fresneda (2) & Arturo Baz (1) (1) Departamento de Zoología y Antropología Física. Facultad de Biología. Universidad de Alcalá. E-28871 Alcalá de Henares, Madrid, Spain (2) Ca de Massa, 25526 Llesp-El Pont de Suert, Lleida, Spain Abstract. A new subspecies of Troglorites breuili Jeannel 1919 (T. breuili salgadoi ssp. n.) which was discovered at Cueva del Viento, Mendaro, Guipúzcoa (Spain), is described. It features a prominent macrocephaly, a strongly transverse pronotum and peculiar cephalic setation. A morphometric analysis is presented, along with a redescription of the nominotypical subspecies —female genitalia are described in detail — and characterization of T. breuili mendizabali Jeannel 1921. The description also includes a chorological update of the three subspecies mentioned above, an inventory of the fauna that lives with each of them, and points are made about their biology and biogeography. Résumé. Nouvelles données sur Troglorites breuili Jeannel 1919 (Coleoptera : Carabidae : Pterostichini) : a hypogean Iberian species with description of a new subspecies. Une nouvelle sous-espèce de Troglorites breuili Jeannel 1919 (T. breuili sagadoi ssp. n.) est décrite de la grotte Cueva del Viento, Mendaro, Guipúzcoa (Espagne). Elle se caractérise par une macrocéphalie proéminente, un pronotum fortement transverse et une cheitotaxie céphalique particulière. Une analyse morphométrique est présentée, ainsi qu’une redescription de la sous-espèce nominale, dont les genitalia sont décrits en détail, de même enfi n que T. breuili mendibazali Jeannel 1921. La description inclu aussi une mise à jour de la chorologie de ces trois sous-espèces, une présentation des espèces qui vivent avec chacune d’elles et quelques points sur leur biologie et de leur biogéographie. Keywords: Underground, taxonomy, biology, biogeography, cave. he ground beetle fauna of the Iberian Peninsula Aphaenops], Duvalius Delarouzée 1859, Hydraphaenops Thas a large number of endemic species. From Jeannel 1926, Paraphaenops Jeannel 1916 and Trechus Serrano (2003) 173 species (37% of endemic ground Clairville 1806, a monospecifi c Molopini as Henrotius beetles and 15 % of all Iberian carabids) are exclusively jordai (Reitter 1914) and three Sphodrini from the subterranean (Jiménez-Valverde & Ortuño 2007), subgenus Antisphodrus Schaufuss 1865. occurring in hypogean and endogeous environments. Troglorites is a genus of Pterostichini which has Troglorites breuili Jeannel 1919 is outstanding traditionally been associated with Cryobius Chaudoir amongst the exclusively hypogean species because 1838 (= Haptoderus sensu Jeannel 1942), and more of historical (history of the Iberian biospeleology), closely to the lineage of Pyreneorites Jeannel 1937, ecological, biogeographical, and now, again, for to which it resembles in its general morphology systematic and taxonomic reasons. and certain characteristics of its aedeagus. However, From the historic point of view T. breuili was nowadays, there are no solid arguments to sustain that relationship, and therefore, its systematic position one of the fi rst described species from the hypogean within the tribe remains somewhat enigmatic. environment in Spain (Jeannel 1919b). It is only From an ecological point of view, T. breuili plays surpassed in antiquity by 7 Trechini species from an important role in the Montes Vascos hypogean the genera: Aphaenops Bonvouloir 1862 [Bonvouloir biocenosis, where no other large hypogean Carabidae original spelling is Aphoenops. Subsequently called are known. It has a relatively wide distribution in Aphaenops by nearly all authors. It is possible that the hypogean environments in the Aralar, Urbasa, Andía, name Aphaenops might still be in use (especially in the XX Entzia mountains and the Macizo de Ernio-Zestoa century), as it appears on the ICZN (2000) articles 23.9 (Vives 1980; Ortuño et al. 1996), where it acts as an and 58. Until a defi nitive solution is found the name important predator of invertebrates. In relation to biogeography, this species along with T. ochsi Fagniez 1921 (from Maritime Alps, France), is E-mail: [email protected] an example of a lineage showing a Pyrenean-Provençal Accepté le 28 mai 2009 centrifugal dispersion (Español & Mateu 1950). 537 V. M. Ortuño, J. Fresneda & A. Baz Table 1. Origin and number of specimens. Abbreviation caves: A, Cueva de Akelar (Navarra); M1, Cueva de Martintxurito-I (Navarra); M2, Cueva de Martintxurito-II (Navarra); B, Cueva Baztarroa (Navarra); Eb, Cueva de Erbeltz (Navarra); Et, Cueva de Etxabe (Navarra); I, Cueva de Iguarán (Álava); M, Cueva Mendikute (Guipúzcoa); Pa, Cueva de Pagoeta (Guipúzcoa); Tx, Cueva de Txorrote (Guipúzcoa); SZ, Cueva de Sagain-Zelaia (Guipúzcoa); Ek, Sima de Ekain (Guipúzcoa); V, Cueva del Viento (Guipúzcoa). AM1M2B EbEtI MPaTxSZEkV T. b. breuili 13 ♂♂ 4 ♂♂ 2 ♂♂ 2 ♂♂ 5 ♂♂ 17 ♀♀ 4 ♀♀ 1 ♀ 1 ♀ 1 ♀ 1 ♀ 7 ♀♀ T. b. mendizabali 1 ♂ 1 ♂ 1 ♂ 1 ♂ 5 ♀♀ 1 ♀ 1 ♀ T. b. salgadoi ssp. n. 2 ♂♂ 3 ♀♀ Th is fact suggests, according to Jeannel (1942), that Material and methods their lucifugous ancestors lived in the Eocene in the Examined specimens Pyrenees-Provençal range of mountains, before the Alpine orogeny. 74 specimens of Troglorites have been examined (appendix) and deposited in collections J. Fresneda (JF), V.M. Ortuño (VO), Finally, T. breuili is very interesting from a taxonomic J.Mª. Salgado (JS), M. Toribio (MT) and Museo Nacional de point of view, because subspecifi c diff erentiation has Ciencias Naturales de Madrid (MNCNM). Five specimens been suggested on the basis of morphometrics (see are attributable to new subspecies, and the type material is the Jeannel 1921a; Español & Mateu 1950; Español 1951, following one: 1966) into T. breuili breuili and T. breuili mendizabali Holotype: 1♂, Cueva del Viento, Mendaro (Guipúzcoa, Spain), 15-07-1998/26-04-1999, J. Fresneda leg. (V. M. Ortuño col.). Jeannel 1921. Moreover, an obvious spatial disjunction Paratypes: 1♀, idem, 03-05-1997, J. Fresneda leg. & col.; 1♂, exists: the fi rst subespecies is located in mountains of idem, 15-07-1998, J. Fresneda leg. & col.; 1♀, idem, 15-07- western Navarra and eastern Álava , while the second 1998/26-04-1999, J. Fresneda leg. (V. M. Ortuño col.); 1♀, one, more northern, is found in mountains in Ernio- idem, 18-07-2000, J. Fresneda leg. & col. Zestoa in Guipúzcoa. Genitalia study We have captured several specimens of Troglorites in recent years from a cave in Guipúzcoa, which cannot Th e aedeagus was studied following current protocol in studies of Carabidae: extraction of the abdomen and dry preparation, be assigned to either of the previously described mounting it on fi ne card or, in other cases, including the subspecies; characteristics of the chaetotaxy from genitalia in a drop of dimethyl hydantoin formaldehyde both the cephalic and pronotal regions allow us to (DMHF) on a transparent acetate sheet attached to the same distinguish them from the existing two subspecies. Th e pin as the specimen. macrocephalia of this new Troglorites and the isolation Th e study of the female genitalia involves a slightly more com- plex procedure (see Carayon 1969; Ortuño et al. 2003): the of T. b. breuili and T. b. mendizabali, have prompted us abdomen is dissected, and the contents are immersed in a satu- to describe it formally and aff ord it subspecies status. rated KOH solution for 12–18 hours. Membranous structures are then stained with Chlorazol-E® black. Th e genital prepara- tion is made in a DMHF drop on a transparent acetate sheet. Table 2. Mean values and Standard deviations of the morphometric variables used to discriminate subspecies. Morphometric analysis: Methods T. b. breuili T. b. mendizabali T. b. salgadoi ssp. n. n = 58 n = 11 n = 5 A total of 74 specimens from various locations in northern Spain mean sd mean sd mean sd were studied (Table 1 and appendix). Eight morphometric variables were measured (see Fig. 1d). All measurements are in HW 2.033 0.106 2.426 0.126 2.726 0.049 millimetres and were log-transformed for analysis. Mean values HEW 2.096 0.104 2.296 0.119 2.486 0.124 and standard deviations are given in Table 2. To reduce the MEW 3.282 0.13 3.518 0.144 3.832 0.169 number of descriptive variables, a principal component analysis MPW 2.47 0.119 2.749 0.127 3.17 0.1 (PCA) was performed on the eight body measurements. Th e mPW _1 1.628 0.075 1.865 0.065 2.198 0.086 analysis revealed one major independent trend of variation HL 1.627 0.079 1.905 0.087 2.18 0.044 across individuals in terms of body size. Scores of individual EL 5.627 0.198 5.86 0.2 6.34 0.296 beetles on this principal component will be used to characterize PL 2.014 0.1 2.125 0.092 2.112 0.097 them in terms of body size. For sexual dimorphism and population variation, we used analysis of variance (ANOVA) for unbalanced designs to test for diff erences between means. 538 New data and new subspecies of Troglorites breuili Figure 1 Head, pronotum and basal region of the elytra of T. breuili. a, T. .b. breuili; b, T. b. mendizabali; c, T. b. salgadoi ssp. n.. (scale: 2 mm); d, Biometric measures: HL, Head length; HW, Head width; MPW, Maximum pronotum width; mPW, Minimum pronotum width; PL, Pronotum length; HEW, Humeral elytra width; MEW, Maximum elytra width; EL, Elytra length. We used Discriminant Analysis to compare taxa because is Th orax (Fig. 1). Pronotum cordiform, wider than long (length a very useful tool (1) for detecting the variables that allow measured in the plan of bilateral symmetry); fore angles acute; discrimination between groups, and (2) for classifying cases hind angles more or less prominent (slightly acute, right or into groups with better than chance accuracy (Tab.
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