Ann. soc. entomol. Fr. (n.s.

), 2010, 46 (34) : 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. Ortuo (1), Javier Fresneda (2) & Arturo Baz (1)
(1)

Departamento de Zoologa y Antropologa Fsica. Facultad de Biologa. Universidad de Alcal. E-28871 Alcal 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, Guipzcoa (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. Rsum. Nouvelles donnes sur Troglorites breuili Jeannel 1919 (Coleoptera : Carabidae : Pterostichini) : a hypogean Iberian species with description of a new subspecies. Une nouvelle sous-espce de Troglorites breuili Jeannel 1919 (T. breuili sagadoi ssp. n.) est dcrite de la grotte Cueva del Viento, Mendaro, Guipzcoa (Espagne). Elle se caractrise par une macrocphalie prominente, un pronotum fortement transverse et une cheitotaxie cphalique particulire. Une analyse morphomtrique est prsente, ainsi quune redescription de la sous-espce nominale, dont les genitalia sont dcrits en dtail, de mme enn que T. breuili mendibazali Jeannel 1921. La description inclu aussi une mise jour de la chorologie de ces trois sous-espces, une prsentation des espces qui vivent avec chacune delles et quelques points sur leur biologie et de leur biogographie.

Keywords: Underground, taxonomy, biology, biogeography, cave.

he ground beetle fauna of the Iberian Peninsula has a large number of endemic species. From Serrano (2003) 173 species (37% of endemic ground beetles and 15 % of all Iberian carabids) are exclusively subterranean (Jimnez-Valverde & Ortuo 2007), occurring in hypogean and endogeous environments. Troglorites breuili Jeannel 1919 is outstanding amongst the exclusively hypogean species because of historical (history of the Iberian biospeleology), ecological, biogeographical, and now, again, for systematic and taxonomic reasons. From the historic point of view T. breuili was one of the rst described species from the hypogean environment in Spain (Jeannel 1919b). It is only surpassed in antiquity by 7 Trechini species from the genera: Aphaenops Bonvouloir 1862 [Bonvouloir original spelling is Aphoenops. Subsequently called Aphaenops by nearly all authors. It is possible that the name Aphaenops might still be in use (especially in the XX century), as it appears on the ICZN (2000) articles 23.9 and 58. Until a denitive solution is found the name

E-mail: vicente.ortuno@uah.es Accept le 28 mai 2009

Aphaenops], Duvalius Delarouze 1859, Hydraphaenops Jeannel 1926, Paraphaenops Jeannel 1916 and Trechus Clairville 1806, a monospecic Molopini as Henrotius jordai (Reitter 1914) and three Sphodrini from the subgenus Antisphodrus Schaufuss 1865. Troglorites is a genus of Pterostichini which has traditionally been associated with Cryobius Chaudoir 1838 (= Haptoderus sensu Jeannel 1942), and more closely to the lineage of Pyreneorites Jeannel 1937, to which it resembles in its general morphology and certain characteristics of its aedeagus. However, nowadays, there are no solid arguments to sustain that relationship, and therefore, its systematic position within the tribe remains somewhat enigmatic. From an ecological point of view, T. breuili plays an important role in the Montes Vascos hypogean biocenosis, where no other large hypogean Carabidae are known. It has a relatively wide distribution in hypogean environments in the Aralar, Urbasa, Anda, Entzia mountains and the Macizo de Ernio-Zestoa (Vives 1980; Ortuo et al. 1996), where it acts as an important predator of invertebrates. In relation to biogeography, this species along with T. ochsi Fagniez 1921 (from Maritime Alps, France), is an example of a lineage showing a Pyrenean-Provenal centrifugal dispersion (Espaol & Mateu 1950).

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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 Iguarn (lava); M, Cueva Mendikute (Guipzcoa); Pa, Cueva de Pagoeta (Guipzcoa); Tx, Cueva de Txorrote (Guipzcoa); SZ, Cueva de Sagain-Zelaia (Guipzcoa); Ek, Sima de Ekain (Guipzcoa); V, Cueva del Viento (Guipzcoa). T. b. breuili T. b. mendizabali T. b. salgadoi ssp. n. A 13 17 M1 4 4 M2 2 1 B 1 Eb 2 1 Et 1 I 5 7 M Pa Tx SZ Ek V

1 5

1 1

1 1 2 3

This fact suggests, according to Jeannel (1942), that their lucifugous ancestors lived in the Eocene in the Pyrenees-Provenal range of mountains, before the Alpine orogeny. Finally, T. breuili is very interesting from a taxonomic point of view, because subspecic dierentiation has been suggested on the basis of morphometrics (see Jeannel 1921a; Espaol & Mateu 1950; Espaol 1951, 1966) into T. breuili breuili and T. breuili mendizabali Jeannel 1921. Moreover, an obvious spatial disjunction exists: the rst subespecies is located in mountains of western Navarra and eastern lava , while the second one, more northern, is found in mountains in ErnioZestoa in Guipzcoa. We have captured several specimens of Troglorites in recent years from a cave in Guipzcoa, which cannot be assigned to either of the previously described subspecies; characteristics of the chaetotaxy from both the cephalic and pronotal regions allow us to distinguish them from the existing two subspecies. The macrocephalia of this new Troglorites and the isolation of T. b. breuili and T. b. mendizabali, have prompted us to describe it formally and aord it subspecies status.
Table 2. Mean values and Standard deviations of the morphometric variables used to discriminate subspecies. T. b. breuili n = 58 mean sd 2.033 0.106 2.096 0.104 3.282 0.13 2.47 0.119 1.628 0.075 1.627 0.079 5.627 0.198 2.014 0.1 T. b. mendizabali n = 11 mean sd 2.426 0.126 2.296 0.119 3.518 0.144 2.749 0.127 1.865 0.065 1.905 0.087 5.86 0.2 2.125 0.092 T. b. salgadoi ssp. n. n=5 mean sd 2.726 0.049 2.486 0.124 3.832 0.169 3.17 0.1 2.198 0.086 2.18 0.044 6.34 0.296 2.112 0.097

Material and methods Examined specimens 74 specimens of Troglorites have been examined (appendix) and deposited in collections J. Fresneda (JF), V.M. Ortuo (VO), J.M. Salgado (JS), M. Toribio (MT) and Museo Nacional de Ciencias Naturales de Madrid (MNCNM). Five specimens are attributable to new subspecies, and the type material is the following one: Holotype: 1, Cueva del Viento, Mendaro (Guipzcoa, Spain), 15-07-1998/26-04-1999, J. Fresneda leg. (V. M. Ortuo col.). Paratypes: 1, idem, 03-05-1997, J. Fresneda leg. & col.; 1, idem, 15-07-1998, J. Fresneda leg. & col.; 1, idem, 15-071998/26-04-1999, J. Fresneda leg. (V. M. Ortuo col.); 1, idem, 18-07-2000, J. Fresneda leg. & col. Genitalia study The aedeagus was studied following current protocol in studies of Carabidae: extraction of the abdomen and dry preparation, mounting it on ne card or, in other cases, including the genitalia in a drop of dimethyl hydantoin formaldehyde (DMHF) on a transparent acetate sheet attached to the same pin as the specimen. The study of the female genitalia involves a slightly more complex procedure (see Carayon 1969; Ortuo et al. 2003): the abdomen is dissected, and the contents are immersed in a saturated KOH solution for 1218 hours. Membranous structures are then stained with Chlorazol-E black. The genital preparation is made in a DMHF drop on a transparent acetate sheet. Morphometric analysis: Methods A total of 74 specimens from various locations in northern Spain were studied (Table 1 and appendix). Eight morphometric variables were measured (see Fig. 1d). All measurements are in millimetres and were log-transformed for analysis. Mean values and standard deviations are given in Table 2. To reduce the number of descriptive variables, a principal component analysis (PCA) was performed on the eight body measurements. The analysis revealed one major independent trend of variation across individuals in terms of body size. Scores of individual beetles on this principal component will be used to characterize them in terms of body size. For sexual dimorphism and population variation, we used analysis of variance (ANOVA) for unbalanced designs to test for dierences between means.

HW HEW MEW MPW mPW _1 HL EL PL

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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 a very useful tool (1) for detecting the variables that allow discrimination between groups, and (2) for classifying cases into groups with better than chance accuracy (Tab. 2).

Taxonomy
Troglorites breuili Jeannel 1919 (Figs. 14)
Redescription
Troglorites breuili Jeannel 1919, Bulletin de la Socit entomologique de France, 1918: 273.

Thorax (Fig. 1). Pronotum cordiform, wider than long (length measured in the plan of bilateral symmetry); fore angles acute; hind angles more or less prominent (slightly acute, right or subright); lateral gutter deep and regularly broad; basal foveae large, smooth and deep; disk divided longitudinally by deep and narrow central furrow, branching o towards anterior and posterior angles. Pronotal chaetotaxy: one posterior seta next to hind angle and two anterior setae on each side (a single seta in some specimens). Metanotum with vestigial wings (squamiform) and

Length: 9.013.2 mm (from tip of mandible to elytron apex). Anophthalmic, discoloured (rufo-testaceous colour) with glabrous integument. Head (Figs. 12). Voluminous head. Prominent ocular convexity with a small ocular scar on front side and a few vestigial discoloured ommatidia. Wide cephalic disc with a very reduced supraocular groove, which features a fronto-clypeal indentation lengthways. Transverse rectangular labrum, with slight notch in apical margin. Prominent, sharp mandible. Maxillary palps without setae and last fusiform palpomere. Labial palps with two setae and last fusiform palpomere. Labium with prominent and bid triangular shaped tooth. Prominent epiloba. Filiform antennae densely setulose from the 4th antennomere. Cephalic chaetotaxy: six setae on labrum, more elongated lateral setae; one seta on both sides of clypeus; two pairs (anterior and posterior) of supraocular setae (some specimens have a supernumerary setigerous pore next to the posterior seta); prebasilar with two pairs of setae (or even three).

Figure 2 Head in ventral view of T. b. breuili from Cueva de Akelar (Navarra) (scale: 1 mm).

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V. M. Ortuo, J. Fresneda & A. Baz

a pair of large spiracles (Fig. 3a). Prosternum with cordiform intercoxal apophysis, this structure without dened marginal furrow. Episternum: proepisternum and mesoepisternum smooth. Metaepisternum very short and smooth. Elytra. Elliptical overall. Epipleures crossed. Humeral region obtuse, with a dened elongated margin up to 1st stria. Basal region depressed (from rear angle up to scutellum). Maximum width of the elytra occurs almost half way along their length. No apical stria. Eight well dened linear striae from the base to the apex (some of which are often anastomosan). Interstrial intervals convex. Elytral chaetotaxy: scutellar setae on 2nd stria, from six to eight discal setae on the 3rd interstrial interval and two or three on the 5th, one apical seta on 7th interstria, 15 umbilical setal pores, distributed into two sets: a fore set including six setae and a rear set including nine setae. Legs: trochanters with no outstanding features. Protibia glabrous. Protibial cleaning organ with two clip setae in internal notch. Mesotibial cleaning organ comb like (sensu Ortuo 1990). Mesotarsus and metatarsus grooved. Onychium glabrous underneath. Claws smooth. The protarsus of male specimens

with rst three tarsomeres expanded and two rows of adhesive phanerae in the middle instead. Abdomen. Males with a smooth fovea in the last sternite; apical margin slightly sinuate (Fig. 3b). Last sternite of males and females (Fig. 3c) with two and four setae respectively. Ring of the genital segment of the male after gure 3d. Male genitalia (Fig. 3e, 3f ). Median lobe with a well developed spatula-shaped apical margin. Parameres asymmetrical, the left one ear-shaped,the right one short and virgulate. Inter and intrapopulational variation occurs in the width and perimeter of the apical margin of the median lobe. Inner sac of the aedeagus with a small sclerotized piece. Female genitalia (Fig. 4). The anatomy of the female genitalia of T. breuili is illustrated in Mateu (1997) but there are some inaccuracies, apart from not being formally described. External genitalia formed by dimerous IX gonopods (gonocoxites and gonosubcoxites) and IX laterotergites. IX gonocoxite unguiform and highly sclerotized, with three thorn-shaped setae of considerable size on dorsal surface (two setae located near external margin); small groove near apex and above ventral

Figure 3 a, Metanotum metatergal apparatus of T. b. breuili from Cueva de Akelar (Navarra); last abdominal sternite of the b, male and c, the female of T. b. breuili from Cueva de Martintxurito 2; d, ring of the genital segment of the male of T. b. breuili from Cueva de Akelar (Navarra); aedeagus e, in rigth lateral view and f, left lateral view of T. b. breuili from Cueva de Akelar (Navarra). (scales, a, e and f: 0.5 mm; b, c and d: 1 mm).

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surface, with two ne sensorial setae (nematiform setae). IX gonosubcoxite subtriangular, slightly longer than wider, with three smaller setae found in areas close to base. IX laterotergite wing-shaped, slightly sclerotized with one group of setae near basal margin (approximately 12). Spermathecal complex largely membranous. Vagina and bursa copulatrix comprised of a large chalice-shaped bag with two sclerotized areas. Spermatheca digitiform. Spermathecal duct long and narrow (4 or 5 times longer than spermatheca) with a small sclerotized area on base, next to oviduct. Spermathecal gland globular, attached to the base of spermatheca by means of a narrow and short duct.

Table 3. Summary of the Discriminant Analysis. Discriminant function 1 2 Eigenvalue 10.3436 1.022 Percent relative 91 9 Canonical correlation 0.955 0.711 P-value 0.000 0.000

Subspecies When Troglorites breuili mendizabali was described, this was done based on its geographic isolation from the typical form, and some quantitative morphologic characters: la cabeza es siempre mucho ms voluminosa; orbicular, tan ancha como la base de los litros. Por consecuencia, el pronoto est mucho ms ensanchado adelante que en los T. breuili tpicos (Jeannel 1921a). The discovery of a new population of this species, distant to the area of distribution of T. b. breuili and T. b. mendizabali, follows the guideline of this last one. These specimens are highly macrocephalous, accompanied

by a remarkable widening of the pronotum (Fig. 1c). The isolation of these new populations, coupled with morphometric data and unique features of the setation of both head and pronotum allow us to recognize these specimens as a third subspecies: Troglorites breuili salgadoi ssp. n. The recognition of subspecies of T. breuili by means of a morphometric study is made for the rst time.

Troglorites breuili salgadoi ssp. n. (Fig. 1c)

Ground beetle of greater dimensions than those of other subspecies (length: 11.513.2 mm). Pronotum more transverse (mean PL/MPW = 0.66) (Fig. 1c) and sinuate near posterior angles that in T. b. mendizabali (mean PL/MPW = 0.77) (Fig. 1b) and T. b. breuili (mean PL/MPW = 0.81) (Fig. 1a). Clearly macrocephalous. In all specimen two signicant qualitative dierences have been observed from T. b. breuili and T. b. mendizabali. The new subspecies tends to have a single anterior marginal setae whereas the most common situation in the other subspecies is to have two or three setae. In addition, in the new subspecies the tendency is to duplicate (rarely to triplicate) the posterior supraocular setigerous pore. Male and female genitalia do not show signicant dierences between subspecies.

Discussion
Morphometry No sexual size dimorphism has been found in Troglorites breuili (ANOVA of PCA scores non-

Figure 4 Morphology of the female genitalia of T. breuili breuili from Cueva de Martintxurito: a, genital shield; b, spermathecal complex in dorsal view and, c, ventral view. (scales, a: 0.25 mm; b and c: 0.5 mm).

Figure 5 Discriminant functions graphs. Cases (individuals) were plotted on the basis of the values for two canonical discriminant functions.

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Table 4. Coecients of the classication functions for each one of the three T. breuili subspecies. T. b. breuili HW HEW MEW MPW mPW _1 HL EL PL CONSTANT 174.081 152.587 155.014 71.714 145.465 167.941 146.261 34.715 461.716 T. b. mendizabali 102.021 156.392 149.748 112.508 192.188 207.298 130.491 56.006 506.17 T. b. salgadoi ssp. n. 140.331 176.482 171.61 66.675 263.12 262.152 137.912 159.825 656.908

signicant F = 0.19, df = 73, P = 0.6671). As a consequence, data from males and females were pooled in subsequent analyses. It is necessary to point out that this analysis assumes that the size dierences in both sexes are similar in all the populations of T. breuili. Troglorites breuili present in the caves by very small number of specimens, making comparisons amongst populations dicult. The three subspecies dier in the size of the body (F = 64.23, df = 73, P = 0.00), a body size gradient being observed from T. b. breuili (the smallest) to T. b. salgadoi ssp. n. (the biggest) (Fig. 5). A discriminant analysis was performed in which 74 cases were used to develop a model to discriminate amongst the 3 subspecies of T. breuili. The two discriminant functions, based on eight variables, are signicant (P < 0.05) (see the summary of the Discriminant Analysis in the Table 3). We can visualize how the two functions discriminate between groups by plotting the individual scores for the two discriminant functions (Fig. 6). Another major purpose to which discriminant analysis is applied is the issue of predictive classication of

cases. The derived classication functions (Table 4) can be used to determine to which group each case most likely belongs. There are as many classication functions as there are groups. Each function allows us to compute classication scores for each case for each group. For example, the classication function for T. b. breuili would be as follows: 461.716 174.081 HW 152.587 HEW 71.714 MPW + 145.465 mPW_1 + 167.941 HL + 146.261 EL 34.715 PL Once we have computed the classication scores for a case, it is easy to decide how to classify the case: in general we classify the case as belonging to the group for which it has the highest classication score. In our case, amongst the 74 observations used to adjust the model, 74 or 100% were classied correctly. That is to say, the three studied taxa can be discriminated without confusion on the basis of a combination of the measured morphometric variables. Chorology Troglorites breuili breuili is extensively distributed around the Sierra de Aralar, at the north of Arakil valley, and in the Sierras de Urbasa, Entzia and Anda, these ones in the south of the valley (Fig. 7). It has been located in several caves which are specically mentioned below. Sierra de Aralar (Navarra): Cueva de Akelar in Alli (type locality, Jeannel 1919b); Cuevas de Martintxurito 1 and 2 in Iribas Jeannel 1919b (it is possible that Cuevas de Ear 1 and 2, also in Iribas, and named after a village inhabited by one of the authors, are another name for the Martintxurito caves); Cueva de Etxabe-Iturri, Cueva de Etxabe and Cueva Baztarroa in Iraeta (Espaol 1948) Cueva de Etxabe-Iturri and Cueva de Etxave are possibly two names for the same cave: although Espaol (1948, 1951) talks of

Figure 6 Plot of the mean of PCA scores, considered as a body size index, for each factor level (subspecies). Bars represent 95% intervals LSD (lowest signicant dierence).

Figure 7 Distribution map of the three subspecies of Troglorites breuili.

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New data and new subspecies of Troglorites breuili

them as if they were two dierent caves in his Iberic pterostquidos compilation; Espaol (1966) only talks about the rst of the two names; Cueva Putxerri near Igaratza refuge in Larraun (Mateu 1945, Espaol 1951) and Cueva de Intzartzu in Ataun (Espaol 1951); Cueva de Basaletz in Huarte-Arakil (Vives 1980). Sierra de Anda (Navarra): Cueva de Erbeltz (Espaol 1948) and Sima Tximua de (Vives 1980) both them in Lizarraga. Sierra de Urbasa (Navarra): Cueva de Larramendikuarro in Larraona and Cueva del Cerro Viejo in Lezaun (Espaol 1951). Sierra de Entzia (lava): Cueva de Iguaran in Salvatierra-Agurain (Ortuo et al. 1996). Troglorites breuili mendizabali lives in mountain ranges which are between the Tolosa, Zestoa and Zarautz, Ernio and Pagoeta massifs, between the Oria and Urola rivers (Fig. 7). It is known in the following caves: Cueva de Ernialde in the village of the same name (type locality, Jeannel 1921a); Cueva de Mendikute (Jeannel 1921a) and Cueva de Txorrote (Espaol 1951), both in Albiztur; Cueva de SagainZelaia in Andazarrate and Sima de Ekain in Zestoa (Vives & Vives 1978); Sima de Itxaropena in Asteasu (Vives 1980); Sima de Alzola in Aia (Galn 2006); and Cueva de Pagoeta in Aia. Troglorites breuili salgadoi ssp. n. is only known in Cueva del Viento, localized in Mendaro (Guipzcoa) (Fig. 7). This cave is situated in the headwaters of the Kilimon river, in the reliefs which are between the Deba and Urola river beds. It is possible that this pterostichine lives in other subterranean spaces in the aforementioned area, like Cueva de Aitbeltz in Andoain or Cueva de Ermitia and Cueva de Arbil in Deba. This hypothesis is based on biogeographical criteria rather than geographical proximity: the troglobitic Leiodidae Quaestus (Quaesticulus) noltei (Coiait 1965) lives in these cave Espaol (1974 which also lives in Cueva del Viento (Salgado et al. 2008) with Troglorites breuili. Biology The rst data on the habitat of T. b. breuili are due to Jeannel (1921a) who indicates: Se encuentra este hermoso carbido andando por el suelo o trepando por las estalagmitas y tratando de esconderse en las grietas. Sus costumbres son ms bien las de un Aphaenops que las de un Ceuthosphodrus. Concluding, in this way: El Troglorites breuili pertenece a la fauna de las paredes estalagmticas, no siendo lapidcola. This armation partly contradicts our observations, and those of other authors (Galn 1993). T. b. breuili is very abundant in Cueva de Akelar and Cuevas de Martintxurito 1 and 2 (= Ear 1 and 2). Here they are always observed as

individuals, walking on the walls, on the limestone stones which cover the cave, or on the oor; however, this hypogean species is also abundant under the clasts deposited on the oor; here, tens of specimens can be found, together with large numbers of E. (Euryspeonomus) breuili (Jeannel 1919) [Coleoptera, Leiodidae]. Nonetheless, T. b. mendizabali is less abundant (personal observation coincidental with Espaol 1951, 1966), although it has been observed and collected under stones in caves. T. b. salgadoi ssp. n. is the rarest subspecies (collected as single individuals in each exploration), and is observed exclusively under clay heaps. In conclusion T. breuili is clearly a typical element of the parietal association in these caves (Jeannel 1921a), however, it can be conrmed (in agreement with other authors (Espaol 1948; Espaol & Mateu 1950; Galn 1993) that this species occupies the hypogean lapidicolous habitat too. It is known that the subsistence of the hypogean environment depends, to a great extent, on epigeous ecosystems (Juberthie & Decu 1994), not only as the original source of the subterranean fauna (from preadapted species Vandel 1964) but especially, by their contribution of energy and materials (and their transporters): it is an environment without solar radiation and for this reason, without autotrophic species (Juberthie & Decu 1994). In consequence, the limited energy resources which arrive from the exterior are quickly degraded by the few members of the extremely short trophic chain. It is possible that Troglorites specimens (highly specialized troglobiomorph facies), are very ecient in the exhaustive exploitation of the limited resources oered by the hypogean environment (especially in deep areas in the caves). It should happen in the subspecies T. b. mendizabali and T. b. salgadoi ssp. n. that have little populations. However, T. b. breuili is the subspecies with a wider distribution and with a bigger population, being very abundant even in the initial section in the caves (places where it can exist a certain inuence of epigeous environment and a bigger material-energy contribution). How can we explain the abundance of such an organism, that occupies the summit of the trophic pyramid? Its abundance may suggest that, unlike the other two subspecies, it is an opportunist organism, with higher adaptive capacity, perhaps in agreement with its probable presence in the more supercial hypogeous environment, the Mesovoid Shallow Substratum (the MSS of Juberthie et al. 1980a, b). This greater amplitude of ecological niche, would explain not only its ample distribution, but also the abundant populations of this predator that could avoid periods of scarcity of food in the caves by moving into the MSS. The fact that this subspecies is able

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Table 5. Coexisting fauna (troglophilic and trogloxenic) of Troglorites breuili. Abbreviation caves: M, Cueva Mendikute (Guipzcoa); Er, Cueva de Ernialde (Guipzcoa); Tx, Cueva de Txorrote (Guipzcoa); SZ, Cueva de Sagain-Zelaia (Guipzcoa); M2, Cueva de Martintxurito-II (Navarra); Ek, Sima de Ekain (Guipzcoa); A, Cueva de Akelar (Navarra); Pu, Cueva de Putxerri (Navarra); B, Cueva Baztarroa (Navarra); Et, Cueva de Etxabe (Navarra); Eb, Cueva de Erbeltz (Navarra); Al, Sima de Alzola (Guipzcoa). TAXA Troglophilic and regular trogloxenic organisms Hirudinea Herpobdellidae Herpobdella octoculata (L. 1758) Gasteropoda Zonitidae Oxychillus arcasianus (Servain 1880) Opiliones Phlangiidae Megabunus diadema (Fabricius 1779) Leiobunum rotundum (Latreille 1798) Ischyropsalidae Ischyropsalis nodifera (Simon 1879) Gyanthidae Gyas titanus (Simon 1879) Travuniidae Peltonychia piochardi (Simon 1892) Araneae Linyphiidae Birgerius microps (Simon 1911) Troglohyphantes furcifer (Simon 1884) Troglohyphantes cantabricus (Simon 1911) Tetragnathidae Meta menardi (Latreille 1804) Meta bourneti Simon 1922 Metellina merianae (Scopoli 1763) Agelenidae Tegenaria inermis Simon 1870 Dictynidae Chorizomma subterraneum Simon 1872 Erigonidae Diplocephalus foraminifer thyrsiger (Simon 1884) Nesticidae Nesticus cellulanus (Clerck 1758) Amaurobidae Amaurobius sp. Isopoda Oniscidae Oniscus asellus L. 1758 Diplopoda Glomeridae Loboglomeris rugifer (Verhoe 1906) Chilopoda Lithobiidae Lithobius pilicornis doriae (Pocock 1890) Lithobius validus vasconicus Chalande 1905 Lithobius tricuspis Meinert 1872 Lithobius romanus inopinatus Matic & Darabantzu 1968 M Er Tx SZ M2 CAVES Ek A Pu B Et Eb Al REFERENCES Galn (2006) + Galn (2006) +

+ +

+ Rambla (1980)

+ + + + +

+ + + + + + + + + + + + Galn (2006) + Galn (2006) + + + + + + + + + Ribera (1980) Galn (2006) Machado (1940)

+ + + +

Serra (1980) Galn (2006)

544

New data and new subspecies of Troglorites breuili

TAXA Troglophilic and regular trogloxenic organisms Diptera Sciaridae Lycosia sp. Mycetophilidae Tarnania fenestralis (Meigen 1818) Culicidae Culex pipiens L. 1758 Trichoptera Limnephilidae Micropterna nycterobia Mac Lachlan 1875 Lepidoptera Geometridae Triphosa dubitata (L. 1758) Coleoptera Carabidae Trechus barnevillei Pandell 1867 Trechus fulvus Dejean 1831 Pterostichus cantabricus heydenianus Jacobson 1907 Laemostenus (Actenipus) oblongus (Dejean 1828) Leiodidae Catops fuliginosus Erichson 1837

Er

Tx

SZ

M2

CAVES Ek A

Pu

Et

Eb

Al

REFERENCES

+ Galn (2006) + + Galn (2006) + Galn (2006) +

+ + + + + + + + + + Vives (1980) Espaol (1948)

to occupy any habitat inside the caves (lapidicolous or parietal) supports this hypothesis. Another hypothesis to explain the dearth of mendizabali and salgadoi ssp. n., is that they do not inhabit the caves, but deep karst sites or MSS, hence they may be highly specialized subspecies to those environments, accessing the caves only when living conditions are favorable (adequate resources available, thermal and hygrometric conditions, etc.). It is likely, but not certain, that the three subspecies have the same ecological preferences. In contrast, the structure of karst does not seem relevant, because wherever they inhabit preferably, they access to the caves as evidenced by the observations. No text has previously summarised data on the fauna which coexist with T. breuili. Although dispersed data exist on other species that have been mentioned from some of the caves where this interesting hypogean Pterostichini has been collected. This information has been compiled and the presence of an appreciable number of species which are not strictly hypogeous is revealed (troglophilic and regular trogloxenic organisms Table 5), of hygrophilous and lucifugous habits that, without a doubt, contribute to the energy enrichment of the caves in where T. breuili lives; they very possibly facilitate its existence, although competitive interactions are possibly established among species (for example with Chilopoda). In addition, T. breuili coexists with other

troglobitic species (Table 6). Small Collembola are frequent in these caves, although it is dicult in some cases to determine if they are exclusively hypogean species. Some troglobitic species are known, like the genera Onychiurus Gervais, 1841 [Onychiuridae], Typhlogastrura Bonet 1930 [Hypogastruridae], Pseudosinella Scher 1897 [Entomobryidae], Arrhopalites Brner 1906 [Sminthuridae] (see Bells 1987). Beetles of the family Leiodidae are remarkable in these environments, by virtue of their specic diversity and the abundance of some populations. Two species of Euryspeonomus Jeannel 1919: E. (E.) breuili breuili Jeannel 1919 in the Cueva de Akelar, Martintxurito 1, Martintxurito 2 (Jeannel 1919b), Etxabe-Iturri, Etxabe, Baztarroa (Espaol 1948), Putxerri (Espaol 1974) and Intzartzu (J. Fresneda: unpublished data); E. (Urbasolus) eloseguii Espaol 1948 in the Cueva de Erbeltz (Espaol 1966). Two species of Speocharidius Jeannel 1919: S. (S.) breuili Jeannel 1919 in the Cueva de Ernialde (Jeannel 1919a), Mendikute (Jeannel 1919a), Txorrote (Bolvar 1921), Pagoeta (Espaol 1974), Sagain-Zelaia (Espaol & Bells 1980) and Sima de Alzola (Galn 2006); S. (S.) vivesi Espaol & Bells 1980 in the Cueva de Sagain-Zelaia (Espaol & Bells 1980). Two species of Josettekia Coiait 1952: J. angelinae Bells & Deliot 1983 in the cueva de Akelar (Bells & Deliot 1983) and Ernialde (Salgado et al. 2008) and J. mendizabali (Bolvar 1921) in the Cueva

545

V. M. Ortuo, J. Fresneda & A. Baz

Table 6. Coexisting fauna (troglobitic) of Troglorites breuili. Abbreviation caves: M, Cueva Mendikute (Guipzcoa); Er, Cueva de Ernialde (Guipzcoa); Tx, Cueva de Txorrote (Guipzcoa); SZ, Cueva de Sagain-Zelaia (Guipzcoa); M2, Cueva de Martintxurito-II (Navarra); Ek, Sima de Ekain (Guipzcoa); A, Cueva de Akelar (Navarra); Pu, Cueva de Putxerri (Navarra); Al, Sima de Alzola (Guipzcoa); M1, Cueva de Martintxurito-I (Navarra). TAXA Troglobitic organisms Oligochaeta Haplotaxidae Haplotaxis navarrensis Delay 1973 Asellota Stenasellidae Stenasellus breuili (Racovitza 1924) Copepoda Ameiridae Stygonitrocrella dubia (Chappuis 1937) Nitocrella vasconica Chappuis 1937 Isopoda - Oniscoidea Trichoniscidae Trichoniscoides pseudomixtus (Arcangeli 1934) Trichoniscoides dubius (Arcangeli 1935) Trichoniscoides cavernicola (Budde-Lund 1885) Pseudoscorpionida Chtoniidae Chthonius (Ephippiochthonius) distinguendus Beier 1930 Neobisiidae Neobisium (Blothrus) nonidezi (Bolvar 1924) Neobisium (Blothrus) breuili (Bolvar 1924) Neobisium (Blothrus) vasconicum (Nonidez 1925) Neobisium sp. Araneae Linyphiidae Troglohyphantes alluaudi Fage 1919 Micrargus cupidon (Simon 1913) Leptyphantes bolivari Fage 1931 Diplopoda Vandeleumidae Vandeleuma hispanica (Ceuca 1967) Julidae Mesoilus cavernarum (Verhoe 1938) Mesoilus henroti (Mauris 1971) Mesoilus stammeri (Verhoe 1936) Chilopoda Lithobiidae Lithobius (Lithobius) anophthalmus Mati 1957 Lithobius (Lithobius) crypticola alavicus Mati 1959 Lithobius (Lithobius) piceus gracilitarsis Brleman 1898 Lithobius (Monotarsobius) reiseri Verhoe 1900 + + + + + + + Serra (1980) Galn (2006) + + + + + + + + + + + + + Ceuca (1967) Vicente (1980) Mauries (1971) Galn (2006) + + Ribera (1980) Bells (1987) Galn (2006) + + + + + + + Estany (1980) + + + + Galn (2006) + + + + + + + + Escol (1980) Arcangeli (1935) Galn (2006) + Margalef (1953) + Magniez (1966) M Er Tx SZ CAVES M2 Ek A Pu Al M1 BIBLIOGRAFA Delay (1973)

546

New data and new subspecies of Troglorites breuili

TAXA Troglobitic organisms Diplura Campodeidae Podocampa simonini Cond 1956 Litocampa espanoli Cond 1950 Coleoptera Pselaphidae Prionobythus bolivari Jeannel 1921 Coleoptera Leiodidae Numerous species

Er

Tx

SZ

CAVES M2 Ek

Pu

Al

M1

BIBLIOGRAFA Cond (1956)

+ +

Galn (2006)

Jeannel (1921b) + In the text +

de Mendikute (Bolvar 1921) and Pagoeta (Espaol 1974). Two species of Bathysciola Jeannel 1910: B. (B.) rugosa (Sharp 1872) in the Cueva de Txorrote (Bolvar 1921); B. (B.) breuili Bolvar 1921 (= sub. B. azuai Bolvar 1921) in the Cueva de Txorrote (Espaol 1974) and the Sima de Alzola (Galn 2006). It is advisable to indicate that we do not know the accompanying fauna of T. breuili in the Cueva de Basaletz, Larramendikuarro, Cerro Viejo, Iguaran, Sima de Tximua and Itxaropena. With respect to the fauna that coexists with T. b. salgadoi ssp. n. little can be said since it is being studied at the present time. We know that it coexists with hypogeous Leiodidae. The underground complex of the Cueva del Viento is opened to the outside by a small gallery, where a strong wind currently circulates, a characteristic that is repeated in all the sectors of the cave which they are of exiguous dimensions. The rst section is arid although it is not rare to observe specimens of Choleva (Choleva) fagniezi brevistylis Jeannel 1923 [Leiodidae, Cholevinae, Cholevini] feeding on numerous corpses of Gastropoda (Salgado et al. 20032004). Past that sector are zones that already present the peculiar characteristics of underground ecosystems (absence of photoperiod, almost constant temperature in this cave between 6 7 C and relative humidity close to saturation) and with frequent formation of stalagmites. T. b. salgadoi n. ssp. has always been found at the end of a great lateral room at the end of the principal gallery of the cave; in that place a ooded zone exists permanently and the surroundings are covered by clay deposits. The rst evidence of the presence of this new subspecies of hypogean Pterostichini was the observation and collecting of a dead specimen (conserved in the type series) in good state of preservation. It is an extremely rare insect in the typical locality, in comparison with

the other two subspecies, that coexists with two Leiodidae [Cholevinae, Leptodirini], also highly adapted to the environment subterranean: Aranzadiella leizaolai Espaol 1972 (Fresneda & Salgado 2000) and Quaestus (Quaesticulus) noltei (Coiait 1965). No other Carabidae have been observed coexisting with T. b. salgadoi ssp. n., and no other large hypogean Carabidae are known of other cavities of the region. Biogeography Some of the biogeographical features of the region where the Cueva del Viento is found the river Deba basin should be highlighted. For Leptodirini (Leiodidae, Cholevinae), this area is the boundary between the two phyletic clusters in the Iberian peninsula; on both river banks near the river basin there is a small overlapping area of these two lines of Leptodirini. Therefore one can nd Aranzadiella leizaolai (phyletic series Speonomus) and Quaestus (Quaestus) noltei (phyletic series Quaestus) living together on both banks of the river. A recent divergence date using molecular data has been suggested for the two colonies of Q. (Q.) noltei living on the two river banks, which are morphologically and anatomically identical (Ribera, com. pers.: data from four mitochondrial genes (2,000 pb of cytochrome B, cytochrome oxidase I, NAD dehydrogenase I, and the ribosomal unit 16S) show 0.75% sequence divergence which mean an isolation of populations 300,000 to 400,000 years BP, assuming a 2.3% divergence every million years for insect mitochondrial genomes Brower 1994). This fact proves that the biogeographical border, namely the river Deba, was formed between 300,000 and 400,000 years ago, isolating populations from both sides. It may not be possible to establish that the same evolutionary processes have applied to Carabidae and Leiodidae. However, both the dispersion and

547

V. M. Ortuo, J. Fresneda & A. Baz

colonization of specimens living in underground ecosystems characterized by strict and constant features in the same geographical area may have taken place side by side, given that they are determined by the same climatic conditions. The fact that Troglorites features a high level of adaptation may be due to a long history of evolution in subterranean environments. Nevertheless, the diversication of subspecies (with stable but not very signicant dierences as in Troglorites) is likely to be much more recent. Probably this phenomenon happened when this animal was already totally adapted to underground life. One hypothese is that Troglorites does not live on the west bank of the river Deba as it arrived when it could no longer be reached. On the one hand the West bank of the river Deba was a potential habitat to be colonized and on the other hand the predators are troglophilic species slightly modied. So T. b. salgadoi ssp. n. seems to have arrived to the Eastern bank some time after a 300,000400,000 year-span, when an impassable biogeographical frontier had already been formed. Troglorites must have spread within the time-span mentioned above and due to climate change, from a still undetermined scattering center. An immediate consequence of the data we have, is the following hypothesis: this colonization could be characterized as a sequence of periods of expansion and settlement into areas of refuge; given that the available time for expansion was limited, the colonizing population seems to have been made up of a reduced number of specimens (founder eect); which might in turn have splitted into the above mentioned subspecies in a relatively short period of time.
Acknowledgements. We would like to make acknowledgements to Jos Mari Rey for providing the data as regards varied place names of underground areas of Navarra; Jos M Salgado and Marcos Toribio for allowing us to borrow Troglorites specimens, Ignacio Ribera for the molecular data about Leptodirini, and Alberto Sendra for validating the information about Diplura of the caves from Navarra. And we wish to thank to David Bilton to read and check the English and suggest many improvements to a rst draft.

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