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 complex 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 preparation 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. 2). subright); lateral gutter deep and regularly broad; basal foveae large, smooth and deep; disk divided longitudinally by deep Taxonomy and narrow central furrow, branching oﬀ 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 Troglorites breuili Jeannel 1919 (Figs. 1–4) specimens). Metanotum with vestigial wings (squamiform) and

Redescription Troglorites breuili Jeannel 1919, Bulletin de la Société entomologique de France, 1918: 273. Length: 9.0–13.2 mm (from tip of mandible to elytron apex). Anophthalmic, discoloured (rufo-testaceous colour) with glabrous integument. Head (Figs. 1–2). 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 biﬁ d 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) Figure 2 of supraocular setae (some specimens have a supernumerary Head in ventral view of T. b. breuili from Cueva de Akelar (Navarra) (scale: setigerous pore next to the posterior seta); prebasilar with two 1 mm). pairs of setae (or even three).

539 V. M. Ortuño, J. Fresneda & A. Baz a pair of large spiracles (Fig. 3a). Prosternum with cordiform with fi rst three tarsomeres expanded and two rows of adhesive intercoxal apophysis, this structure without defi ned marginal phanerae in the middle instead. furrow. Episternum: proepisternum and mesoepisternum Abdomen. Males with a smooth fovea in the last sternite; apical smooth. Metaepisternum very short and smooth. margin slightly sinuate (Fig. 3b). Last sternite of males and Elytra. Elliptical overall. Epipleures crossed. Humeral region females (Fig. 3c) with two and four setae respectively. Ring of obtuse, with a defi ned elongated margin up to 1st stria. Basal the genital segment of the male after fi gure 3d. region depressed (from rear angle up to scutellum). Maximum Male genitalia (Fig. 3e, 3f). Median lobe with a well developed width of the elytra occurs almost half way along their length. spatula-shaped apical margin. Parameres asymmetrical, the left No apical stria. Eight well defi ned linear striae from the base one ear-shaped,the right one short and virgulate. Inter and to the apex (some of which are often anastomosan). Interstrial intrapopulational variation occurs in the width and perimeter of intervals convex. Elytral chaetotaxy: scutellar setae on 2nd stria, the apical margin of the median lobe. Inner sac of the aedeagus from six to eight discal setae on the 3rd interstrial interval with a small sclerotized piece. and two or three on the 5th, one apical seta on 7th interstria, Female genitalia (Fig. 4). Th e anatomy of the female genitalia 15 umbilical setal pores, distributed into two sets: a fore set of T. breuili is illustrated in Mateu (1997) but there are some including six setae and a rear set including nine setae. Legs: inaccuracies, apart from not being formally described. External trochanters with no outstanding features. Protibia glabrous. genitalia formed by dimerous IX gonopods (gonocoxites Protibial cleaning organ with two clip setae in internal notch. and gonosubcoxites) and IX laterotergites. IX gonocoxite Mesotibial cleaning organ comb like (sensu Ortuño 1990). unguiform and highly sclerotized, with three thorn-shaped Mesotarsus and metatarsus grooved. Onychium glabrous setae of considerable size on dorsal surface (two setae located underneath. Claws smooth. Th e protarsus of male specimens 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).

540 New data and new subspecies of Troglorites breuili surface, with two fi ne sensorial setae (nematiform setae). IX Table 3. Summary of the Discriminant Analysis. gonosubcoxite subtriangular, slightly longer than wider, with three smaller setae found in areas close to base. IX laterotergite Discriminant Eigenvalue Percent Canonical P-value wing-shaped, slightly sclerotized with one group of setae near function relative correlation basal margin (approximately 12). Spermathecal complex largely 1 10.3436 91 0.955 0.000 membranous. Vagina and bursa copulatrix comprised of a large 2 1.022 9 0.711 0.000 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 by a remarkable widening of the pronotum (Fig. 1c). base of spermatheca by means of a narrow and short duct. Th e isolation of these new populations, coupled with Subspecies morphometric data and unique features of the setation of both head and pronotum allow us to recognize When Troglorites breuili mendizabali was described, these specimens as a third subspecies: Troglorites breuili this was done based on its geographic isolation from salgadoi ssp. n. Th e recognition of subspecies of T. the typical form, and some quantitative morphologic breuili by means of a morphometric study is made for characters: “la cabeza es siempre mucho más voluminosa; the fi rst time. orbicular, tan ancha como la base de los élitros. Por consecuencia, el pronoto está mucho más ensanchado Troglorites breuili salgadoi ssp. n. (Fig. 1c) adelante que en los T. breuili típicos” (Jeannel 1921a). Th e discovery of a new population of this species, Ground beetle of greater dimensions than those of other distant to the area of distribution of T. b. breuili and T. b. subspecies (length: 11.5–13.2 mm). Pronotum more transverse mendizabali, follows the guideline of this last one. Th ese (mean PL/MPW = 0.66) (Fig. 1c) and sinuate near posterior angles that in T. b. mendizabali (mean PL/MPW = 0.77) (Fig. specimens are highly macrocephalous, accompanied 1b) and T. b. breuili (mean PL/MPW = 0.81) (Fig. 1a). Clearly macrocephalous. In all specimen two signifi cant qualitative diff erences have been observed from T. b. breuili and T. b. mendizabali. Th e 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 signifi cant diff erences 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 Figure 5 Martintxurito: a, genital shield; b, spermathecal complex in dorsal view Discriminant functions graphs. Cases (individuals) were plotted on the and, c, ventral view. (scales, a: 0.25 mm; b and c: 0.5 mm). basis of the values for two canonical discriminant functions.

541 V. M. Ortuño, J. Fresneda & A. Baz

Table 4. Coeffi cients of the classifi cation functions for each one of the three cases. Th e derived classifi cation functions (Table 4) T. breuili subspecies. can be used to determine to which group each case T. b. T. b. salgadoi most likely belongs. Th ere are as many classifi cation T. b. breuili mendizabali ssp. n. functions as there are groups. Each function allows us HW –174.081 –102.021 –140.331 to compute classifi cation scores for each case for each HEW –152.587 –156.392 –176.482 group. For example, the classifi cation function for T. b. MEW 155.014 149.748 171.61 breuili would be as follows: MPW –71.714 –112.508 –66.675 – 461.716 – 174.081 HW – 152.587 HEW – mPW _1 145.465 192.188 263.12 71.714 MPW + 145.465 mPW_1 + 167.941 HL + HL 167.941 207.298 262.152 146.261 EL – 34.715 PL EL 146.261 130.491 137.912 Once we have computed the classifi cation scores PL –34.715 –56.006 –159.825 for a case, it is easy to decide how to classify the case: CONSTANT –461.716 –506.17 –656.908 in general we classify the case as belonging to the group for which it has the highest classifi cation score. In our case, amongst the 74 observations used to adjust the signifi cant –F = 0.19, df = 73, P = 0.6671). As a model, 74 or 100% were classifi ed correctly. Th at is consequence, data from males and females were pooled to say, the three studied taxa can be discriminated in subsequent analyses. It is necessary to point out that without confusion on the basis of a combination of this analysis assumes that the size diff erences in both the measured morphometric variables. sexes are similar in all the populations of T. breuili. Chorology Troglorites breuili present in the caves by very small number of specimens, making comparisons amongst Troglorites breuili breuili is extensively distributed populations diffi cult. Th e three subspecies diff er in the around the Sierra de Aralar, at the north of Arakil size of the body (F = 64.23, df = 73, P = 0.00), a body valley, and in the Sierras de Urbasa, Entzia and Andía, size gradient being observed from T. b. breuili (the these ones in the south of the valley (Fig. 7). It has been located in several caves which are specifi cally smallest) to T. b. salgadoi ssp. n. (the biggest) (Fig. 5). A mentioned below. Sierra de Aralar (Navarra): Cueva de discriminant analysis was performed in which 74 cases Akelar in Alli (type locality, Jeannel 1919b); Cuevas de were used to develop a model to discriminate amongst Martintxurito 1 and 2 in Iribas –Jeannel 1919b– (it is the 3 subspecies of T. breuili. Th e two discriminant possible that Cuevas de Ear 1 and 2, also in Iribas, and functions, based on eight variables, are signifi cant (P < named after a village inhabited by one of the authors, 0.05) (see the summary of the Discriminant Analysis are another name for the Martintxurito caves); Cueva in the Table 3). We can visualize how the two functions de Etxabe-Iturri, Cueva de Etxabe and Cueva Baztarroa discriminate between groups by plotting the individual in Irañeta (Español 1948) —Cueva de Etxabe-Iturri scores for the two discriminant functions (Fig. 6). and Cueva de Etxave are possibly two names for the Another major purpose to which discriminant analysis same cave: although Español (1948, 1951) talks of is applied is the issue of predictive classifi cation 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 Figure 7 signiﬁ cant diﬀ erence). Distribution map of the three subspecies of Troglorites breuili.

542 New data and new subspecies of Troglorites breuili them as if they were two diff erent caves in his Iberic individuals, walking on the walls, on the limestone “pterostíquidos” compilation; Español (1966) only stones which cover the cave, or on the fl oor; however, talks about the fi rst of the two names—; Cueva Putxerri this hypogean species is also abundant under the clasts near Igaratza refuge in Larraun (Mateu 1945, Español deposited on the fl oor; here, tens of specimens can 1951) and Cueva de Intzartzu in Ataun (Español be found, together with large numbers of E. (Eurys- 1951); Cueva de Basaletz in Huarte-Arakil (Vives peonomus) breuili (Jeannel 1919) [Coleoptera, Leiodi- 1980). Sierra de Andía (Navarra): Cueva de Erbeltz dae]. Nonetheless, T. b. mendizabali is less abundant (Español 1948) and Sima Tximua de (Vives 1980) (personal observation coincidental with Español 1951, both them in Lizarraga. Sierra de Urbasa (Navarra): 1966), although it has been observed and collected Cueva de Larramendikuarro in Larraona and Cueva under stones in caves. T. b. salgadoi ssp. n. is the rar- del Cerro Viejo in Lezaun (Español 1951). Sierra de est subspecies (collected as single individuals in each Entzia (Álava): Cueva de Iguaran in Salvatierra-Agurain exploration), and is observed exclusively under clay (Ortuño et al. 1996). heaps. In conclusion T. breuili is clearly a typical ele- Troglorites breuili mendizabali lives in mountain ment of the parietal association in these caves (Jeannel ranges which are between the Tolosa, Zestoa and 1921a), however, it can be confi rmed (in agreement Zarautz, Ernio and Pagoeta massifs, between the with other authors (Español 1948; Español & Mateu Oria and Urola rivers (Fig. 7). It is known in the 1950; Galán 1993) that this species occupies the hypo- following caves: Cueva de Ernialde in the village of gean lapidicolous habitat too. the same name (type locality, Jeannel 1921a); Cueva It is known that the subsistence of the hypogean de Mendikute (Jeannel 1921a) and Cueva de Txorrote environment depends, to a great extent, on epigeous (Español 1951), both in Albiztur; Cueva de Sagain- ecosystems (Juberthie & Decu 1994), not only as the Zelaia in Andazarrate and Sima de Ekain in Zestoa original source of the subterranean fauna (from prea- (Vives & Vives 1978); Sima de Itxaropena in Asteasu dapted species –Vandel 1964–) but especially, by their (Vives 1980); Sima de Alzola in Aia (Galán 2006); and contribution of energy and materials (and their trans- Cueva de Pagoeta in Aia. porters): it is an environment without solar radiation Troglorites breuili salgadoi ssp. n. is only known in and for this reason, without autotrophic species (Ju- Cueva del Viento, localized in Mendaro (Guipúzcoa) berthie & Decu 1994). In consequence, the limited (Fig. 7). Th is cave is situated in the headwaters of energy resources which arrive from the exterior are the Kilimon river, in the reliefs which are between quickly degraded by the few members of the extremely the Deba and Urola river beds. It is possible that this short trophic chain. It is possible that Troglorites speci- pterostichine lives in other subterranean spaces in the mens (highly specialized troglobiomorph facies), are aforementioned area, like Cueva de Aitbeltz in Andoain very effi cient in the exhaustive exploitation of the lim- or Cueva de Ermitia and Cueva de Arbil in Deba. Th is ited resources off ered by the hypogean environment hypothesis is based on biogeographical criteria rather (especially in deep areas in the caves). It should hap- than geographical proximity: the troglobitic Leiodidae pen in the subspecies T. b. mendizabali and T. b. salga- Quaestus (Quaesticulus) noltei (Coiff ait 1965) lives doi ssp. n. that have little populations. However, T. b. in these cave –Español (1974– which also lives in breuili is the subspecies with a wider distribution and Cueva del Viento (Salgado et al. 2008) with Troglorites with a bigger population, being very abundant even breuili. in the initial section in the caves (places where it can exist a certain infl uence of epigeous environment and Biology a bigger material-energy contribution). How can we Th e fi rst data on the habitat of T. b. breuili are due explain the abundance of such an organism, that occu- to Jeannel (1921a) who indicates: “Se encuentra este pies the summit of the trophic pyramid? Its abundance hermoso carábido andando por el suelo o trepando por may suggest that, unlike the other two subspecies, it is las estalagmitas y tratando de esconderse en las grietas. an opportunist organism, with higher adaptive capac- Sus costumbres son más bien las de un Aphaenops que ity, perhaps in agreement with its probable presence in las de un Ceuthosphodrus”. Concluding, in this way: the more superfi cial hypogeous environment, the Mes- “El Troglorites breuili pertenece a la fauna de las paredes ovoid Shallow Substratum (the MSS of Juberthie et al. estalagmíticas, no siendo lapidícola”. Th is affi rmation 1980a, b). Th is greater amplitude of ecological niche, partly contradicts our observations, and those of other would explain not only its ample distribution, but also authors (Galán 1993). T. b. breuili is very abundant in the abundant populations of this predator that could Cueva de Akelar and Cuevas de Martintxurito 1 and avoid periods of scarcity of food in the caves by mov- 2 (= Ear 1 and 2). Here they are always observed as ing into the MSS. Th e fact that this subspecies is able

543 V. M. Ortuño, J. Fresneda & A. Baz

Table 5. Coexisting fauna (troglophilic and trogloxenic) of Troglorites breuili. Abbreviation caves: M, Cueva Mendikute (Guipúzcoa); Er, Cueva de Ernialde (Guipúzcoa); Tx, Cueva de Txorrote (Guipúzcoa); SZ, Cueva de Sagain-Zelaia (Guipúzcoa); M2, Cueva de Martintxurito-II (Navarra); Ek, Sima de Ekain (Guipúzcoa); 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 (Guipúzcoa).

TAXA CAVES Troglophilic and regular trogloxenic organisms M Er Tx SZ M2 Ek A Pu B Et Eb Al REFERENCES Hirudinea Herpobdellidae Galán (2006) Herpobdella octoculata (L. 1758) + Gasteropoda Zonitidae Galán (2006) Oxychillus arcasianus (Servain 1880) + Opiliones Phlangiidae Megabunus diadema (Fabricius 1779) + + Leiobunum rotundum (Latreille 1798) + Ischyropsalidae Rambla (1980) 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 Ribera (1980) Meta menardi (Latreille 1804) + + Galán (2006) Meta bourneti Simon 1922 + Machado (1940) 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 Galán (2006) Oniscus asellus L. 1758 + Diplopoda Glomeridae Galán (2006) Loboglomeris rugifer (Verhoeﬀ 1906) + Chilopoda Lithobiidae Lithobius pilicornis doriae (Pocock 1890) + Serra (1980) Lithobius validus vasconicus Chalande 1905 + Galán (2006) Lithobius tricuspis Meinert 1872 + Lithobius romanus inopinatus Matic & Darabantzu + 1968

544 New data and new subspecies of Troglorites breuili

TAXA CAVES Troglophilic and regular trogloxenic organisms M Er Tx SZ M2 Ek A Pu B Et Eb Al REFERENCES Diptera Sciaridae Lycosia sp. + Mycetophilidae Galán (2006) Tarnania fenestralis (Meigen 1818) + Culicidae Culex pipiens L. 1758 + Trichoptera Limnephilidae Galán (2006) Micropterna nycterobia Mac Lachlan 1875 + Lepidoptera Geometridae Galán (2006) Triphosa dubitata (L. 1758) + Coleoptera Carabidae Trechus barnevillei Pandellé 1867 + Trechus fulvus Dejean 1831 + + Vives (1980) Pterostichus cantabricus heydenianus Jacobson 1907 + ++ Español (1948) Laemostenus (Actenipus) oblongus (Dejean 1828) + ++ Leiodidae Catops fuliginosus Erichson 1837 +

to occupy any habitat inside the caves (lapidicolous or troglobitic species (Table 6). Small Collembola are parietal) supports this hypothesis. Another hypothesis frequent in these caves, although it is diffi cult in some to explain the dearth of mendizabali and salgadoi ssp. cases to determine if they are exclusively hypogean n., is that they do not inhabit the caves, but deep karst species. Some troglobitic species are known, like the sites or MSS, hence they may be highly specialized genera Onychiurus Gervais, 1841 [Onychiuridae], subspecies to those environments, accessing the caves Typhlogastrura Bonet 1930 [Hypogastruridae], only when living conditions are favorable (adequate re- Pseudosinella Schäff er 1897 [Entomobryidae], sources available, thermal and hygrometric conditions, Arrhopalites Börner 1906 [Sminthuridae] (see Bellés etc.). It is likely, but not certain, that the three subspe- 1987). Beetles of the family Leiodidae are remarkable in cies have the same ecological preferences. In contrast, these environments, by virtue of their specifi c diversity the structure of karst does not seem relevant, because and the abundance of some populations. Two species wherever they inhabit preferably, they access to the of Euryspeonomus Jeannel 1919: E. (E.) breuili breuili caves as evidenced by the observations. Jeannel 1919 in the Cueva de Akelar, Martintxurito No text has previously summarised data on the 1, Martintxurito 2 (Jeannel 1919b), Etxabe-Iturri, fauna which coexist with T. breuili. Although dispersed Etxabe, Baztarroa (Español 1948), Putxerri (Español data exist on other species that have been mentioned 1974) and Intzartzu (J. Fresneda: unpublished data); from some of the caves where this interesting hypogean E. (Urbasolus) eloseguii Español 1948 in the Cueva de Pterostichini has been collected. Th is information has Erbeltz (Español 1966). Two species of Speocharidius been compiled and the presence of an appreciable Jeannel 1919: S. (S.) breuili Jeannel 1919 in the Cueva number of species which are not strictly hypogeous is de Ernialde (Jeannel 1919a), Mendikute (Jeannel revealed (troglophilic and regular trogloxenic organisms 1919a), Txorrote (Bolívar 1921), Pagoeta (Español –Table 5–), of hygrophilous and lucifugous habits that, 1974), Sagain-Zelaia (Español & Bellés 1980) and without a doubt, contribute to the energy enrichment Sima de Alzola (Galán 2006); S. (S.) vivesi Español & of the caves in where T. breuili lives; they very Bellés 1980 in the Cueva de Sagain-Zelaia (Español & possibly facilitate its existence, although competitive Bellés 1980). Two species of Josettekia Coiff ait 1952: interactions are possibly established among species (for J. angelinae Bellés & Deliot 1983 in the cueva de Akelar example with Chilopoda). (Bellés & Deliot 1983) and Ernialde (Salgado et al. In addition, T. breuili coexists with other 2008) and J. mendizabali (Bolívar 1921) in the Cueva

545 V. M. Ortuño, J. Fresneda & A. Baz

Table 6. Coexisting fauna (troglobitic) of Troglorites breuili. Abbreviation caves: M, Cueva Mendikute (Guipúzcoa); Er, Cueva de Ernialde (Guipúzcoa); Tx, Cueva de Txorrote (Guipúzcoa); SZ, Cueva de Sagain-Zelaia (Guipúzcoa); M2, Cueva de Martintxurito-II (Navarra); Ek, Sima de Ekain (Guipúzcoa); A, Cueva de Akelar (Navarra); Pu, Cueva de Putxerri (Navarra); Al, Sima de Alzola (Guipúzcoa); M1, Cueva de Martintxurito-I (Navarra).

TAXA CAVES Troglobitic organisms M Er Tx SZ M2 Ek A Pu Al M1 BIBLIOGRAFÍA Oligochaeta Haplotaxidae Delay (1973) Haplotaxis navarrensis Delay 1973 + Asellota Stenasellidae Magniez (1966) Stenasellus breuili (Racovitza 1924) + Copepoda Ameiridae Margalef (1953) Stygonitrocrella dubia (Chappuis 1937) + Nitocrella vasconica Chappuis 1937 + Isopoda - Oniscoidea Escolà (1980) Trichoniscidae Arcangeli (1935) Trichoniscoides pseudomixtus (Arcangeli 1934) ++ + Galán (2006) Trichoniscoides dubius (Arcangeli 1935) + Trichoniscoides cavernicola (Budde-Lund 1885) + + Pseudoscorpionida Chtoniidae Chthonius (Ephippiochthonius) distinguendus Beier + 1930 Estany (1980) Neobisiidae ++ Galán (2006) Neobisium (Blothrus) nonidezi (Bolívar 1924) + + + Neobisium (Blothrus) breuili (Bolívar 1924) ++++ Neobisium (Blothrus) vasconicum (Nonidez 1925) + Neobisium sp. Araneae Linyphiidae Ribera (1980) Troglohyphantes alluaudi Fage 1919 + ++Bellés (1987) Micrargus cupidon (Simon 1913) ++ Galán (2006) Leptyphantes bolivari Fage 1931 + + Diplopoda Vandeleumidae Ceuca (1967) Vandeleuma hispanica (Ceuca 1967) + + + Vicente (1980) Julidae Mauries (1971) Mesoilus cavernarum (Verhoeff 1938) + + Galán (2006) Mesoilus henroti (Mauriès 1971) + Mesoilus stammeri (Verhoeff 1936) + + Chilopoda Lithobiidae Lithobius (Lithobius) anophthalmus Matič 1957 ++ + + Serra (1980) Lithobius (Lithobius) crypticola alavicus Matič 1959 + Galán (2006) Lithobius (Lithobius) piceus gracilitarsis Bröleman + 1898 + Lithobius (Monotarsobius) reiseri Verhoeff 1900

546 New data and new subspecies of Troglorites breuili

TAXA CAVES Troglobitic organisms M Er Tx SZ M2 Ek A Pu Al M1 BIBLIOGRAFÍA Diplura Campodeidae Condé (1956) Podocampa simonini Condé 1956 ++ Galán (2006) Litocampa espanoli Condé 1950 + Coleoptera Pselaphidae Jeannel (1921b) Prionobythus bolivari Jeannel 1921 + + Coleoptera Leiodidae In the text Numerous species

de Mendikute (Bolívar 1921) and Pagoeta (Español the other two subspecies, that coexists with two 1974). Two species of Bathysciola Jeannel 1910: B. (B.) Leiodidae [Cholevinae, Leptodirini], also highly rugosa (Sharp 1872) in the Cueva de Txorrote (Bolívar adapted to the environment subterranean: Aranzadiella 1921); B. (B.) breuili Bolívar 1921 (= sub. B. azuai leizaolai Español 1972 (Fresneda & Salgado 2000) Bolívar 1921) in the Cueva de Txorrote (Español and Quaestus (Quaesticulus) noltei (Coiff ait 1965). No 1974) and the Sima de Alzola (Galán 2006). other Carabidae have been observed coexisting with It is advisable to indicate that we do not know T. b. salgadoi ssp. n., and no other large hypogean the accompanying fauna of T. breuili in the Cueva Carabidae are known of other cavities of the region. de Basaletz, Larramendikuarro, Cerro Viejo, Iguaran, Sima de Tximua and Itxaropena. Biogeography With respect to the fauna that coexists with T. Some of the biogeographical features of the region b. salgadoi ssp. n. little can be said since it is being where the Cueva del Viento is found –the river studied at the present time. We know that it coexists Deba basin– should be highlighted. For Leptodirini with hypogeous Leiodidae. Th e underground complex (Leiodidae, Cholevinae), this area is the boundary of the Cueva del Viento is opened to the outside by a between the two phyletic clusters in the Iberian small gallery, where a strong wind currently circulates, peninsula; on both river banks near the river basin a characteristic that is repeated in all the sectors of there is a small overlapping area of these two lines of the cave which they are of exiguous dimensions. Th e Leptodirini. fi rst section is arid although it is not rare to observe Th erefore one can fi nd Aranzadiella leizaolai specimens of Choleva (Choleva) fagniezi brevistylis (phyletic series Speonomus) and Quaestus (Quaestus) noltei (phyletic series Quaestus) living together on Jeannel 1923 [Leiodidae, Cholevinae, Cholevini] both banks of the river. A recent divergence date feeding on numerous corpses of Gastropoda (Salgado et using molecular data has been suggested for the two al. 2003–2004). Past that sector are zones that already colonies of Q. (Q.) noltei living on the two river banks, present the peculiar characteristics of underground which are morphologically and anatomically identical ecosystems (absence of photoperiod, almost constant (Ribera, com. pers.: data from four mitochondrial genes temperature –in this cave between 6 °–7 °C– and (2,000 pb of cytochrome B, cytochrome oxidase I, relative humidity close to saturation) and with NAD dehydrogenase I, and the ribosomal unit 16S) frequent formation of stalagmites. T. b. salgadoi n. show 0.75% sequence divergence which mean an ssp. has always been found at the end of a great lateral isolation of populations 300,000 to 400,000 years BP, room at the end of the principal gallery of the cave; assuming a 2.3% divergence every million years for in that place a fl ooded zone exists permanently and insect mitochondrial genomes – Brower 1994). Th is the surroundings are covered by clay deposits. Th e fact proves that the biogeographical border, namely the fi rst evidence of the presence of this new subspecies river Deba, was formed between 300,000 and 400,000 of hypogean Pterostichini was the observation and years ago, isolating populations from both sides. collecting of a dead specimen (conserved in the type It may not be possible to establish that the same series) in good state of preservation. It is an extremely evolutionary processes have applied to Carabidae rare insect in the typical locality, in comparison with and Leiodidae. However, both the dispersion and

547 V. M. Ortuño, J. Fresneda & A. Baz colonization of specimens living in underground Brower A. V. Z. 1994. Rapid morphological radiation and convergence ecosystems characterized by strict and constant features among races of the butterfl y Heliconius erato inferred from patterns of mitochondrial DNA evolution. Proceedings of the National Academy of in the same geographical area may have taken place Sciences, USA 91: 6491-6495. side by side, given that they are determined by the Carayon J. 1969. La spermathèque et les voies génitales femelles des same climatic conditions. Lygaeidés Oxycareninae (Heteroptera). Revue Française d’Entomologie Th e fact that Troglorites features a high level of 31: 196-218. adaptation may be due to a long history of evolution Ceuca T. 1967. Contributions à la connaissance de la faune des diplopodes in subterranean environments. Nevertheless, the cavernicoles d’Espagne. Publicaciones del Instituto de Biología Aplicada, Barcelona 42: 127-132. diversifi cation of subspecies (with stable but not Condé B. 1956. Matériaux pour une Monographie des Diploures very signifi cant diff erences as in Troglorites) is likely Campodéidés. Mémoires du Muséum d’Histoire Naturelle. Paris, s. A. to be much more recent. Probably this phenomenon Zool. 12: 1-202. happened when this animal was already totally adapted Delay B. 1973. Deux nouveaux Oligochètes Haplotaxidae troglobies de to underground life. l’Espagne: Haplotaxis navarrensis n. sp. et Haplotaxis cantabronensis n. One hypothese is that Troglorites does not live on the sp. Annales de Spéléologie 28: 405-411. Escolà O. 1980. Crustacea, p. 539-541 in: Bellés X., Blas M., Comas west bank of the river Deba as it arrived when it could J., Escolá O., Español F., Estany J., Rambla R., Ribera C., Serra no longer be reached. On the one hand the West bank A., Vicente M. C., Vives E. Contribución al conocimiento de la fauna of the river Deba was a potential habitat to be colonized cavernícola del País Vasco. Kobie (Bilbao) 10: 525-568. and on the other hand the predators are troglophilic Español F. 1948. Resultados de una campaña bioespeleológica realizada en species slightly modifi ed. So T. b. salgadoi ssp. n. seems la región media occidental de Navarra durante la segunda quincena de to have arrived to the Eastern bank some time after septiembre de 1947. Coleópteros. Eos 24: 233-245. Español F. 1951. Los Pterostichidae hipogeos del Pirineo español a 300,000–400,000 year-span, when an impassable (Coleoptera Adephaga). I Congreso Internacional del Pirineo. Instituto biogeographical frontier had already been formed. de Estudios Pirenaicos, pp: 5-19. Troglorites must have spread within the time-span Español F. 1966. Los pterostíquidos cavernícolas de la Península Ibérica e mentioned above and due to climate change, from a Islas Baleares (col. Caraboidea). Publicaciones del Instituto de Biología still undetermined scattering center. An immediate Aplicada, Barcelona 41: 49-68. consequence of the data we have, is the following Español F. 1974. Los Bathysciinae cavernícolas de Vizcaya, Guipúzcoa y vecinos relieves navarros. Col. Catopidae. Kobie 5: 7-16. hypothesis: this colonization could be characterized Español F., Bellés X. 1980. Coleoptera, Catopidae, Bathysciinae, p. as a sequence of periods of expansion and settlement 557-561 in: Bellés X., Blas M., Comas J., Escolá O., Español F., into areas of refuge; given that the available time for Estany J., Rambla R., Ribera C., Serra A., Vicente M. C., Vives E. expansion was limited, the colonizing population Contribución al conocimiento de la fauna cavernícola del País Vasco. seems to have been made up of a reduced number of Kobie (Bilbao) 10: 525-568. specimens (founder eff ect); which might in turn have Español F., Mateu M. 1950. Sobre algunos insectos cavernícolas del país vasco-navarro. Munibe. Suplemento de Ciencias Naturales del Boletín de splitted into the above mentioned subspecies in a la Real Sociedad Vascongada de los Amigos del País 2: 177-183. relatively short period of time. Estany J. 1980. Arácnida, Pseudoescorpionida, p. 526-528 in: Bellés Acknowledgements. We would like to make acknowledgements X., Blas M., Comas J., Escolá O., Español F., Estany J., Rambla to José Mari Rey for providing the data as regards varied place R., Ribera C., Serra A., Vicente M. C., Vives E. Contribución al names of underground areas of Navarra; José Mª Salgado and conocimiento de la fauna cavernícola del País Vasco. Kobie (Bilbao) 10: Marcos Toribio for allowing us to borrow Troglorites specimens, 525-568. Ignacio Ribera for the molecular data about Leptodirini, and Fresneda J., Salgado J. M. 2000. Revisión de los géneros de Leptodirinae Alberto Sendra for validating the information about Diplura of de la sección Speonomus del sur de Pirineos, II: géneros Aranzadiella the caves from Navarra. And we wish to thank to David Bilton Español 1972, Euryspeonomus Jeannel 1919, Kobiella Español & to read and check the English and suggest many improvements Bellés 1980 y Speocharidius Jeannel 1919. (Coleoptera, Cholevidae). to a fi rst draft. Mémoires de Biospéologie 27: 41-52. Galán C. 1993. Fauna hipogea de Guipúzcoa: su ecología, biogeografía y References evolución. Munibe, número monográfi co, 3-163. Arcangeli A. 1935. Isopodi terrestre di caverne della Spagna (Collezione Galán C. 2006. Fauna cavernícola y poblaciones bacteriales de la sima y río del Museo di Storia naturale di Madrid). Eos 10: 171-195. subterráneo de mondmilch de Alzola (Gipuzkoa). Cota Cero, Periódico Bellés X. 1987. Fauna cavernícola i intersticial de la Península Ibèrica i les digital del mundo subterráneo. Año 2. Número 15, 13 pp. Illes Balears. CSIC-Ed. Moll, Mallorca, 207 p. ICZN 2000. International Commission on Zoological Nomenclature. In- Bellés X., Deliot P. 1983. Nouveaux Bathysciinae (Coléoptères Catopidae) ternational Code of Zoological Nomenclature. Fourth edition. Internatio- des Pyrénées francaises et espagnoles. Mémoires de Biospéologie 10: nal Trust for Zoological Nomenclature. London. xxix + 306 p. 235-242. Jeannel R. 1919a. Bathysciinae nouveaux des Pyrénées espagnoles. Boletín Bolívar C. 1921. Silphidae, p. 524-539 in: Bolívar y Pieltain C., Jeannel de la Sociedad Española de Historia Natural 19: 129-137. R. Coleópteros cavernícolas nuevos de las provincias vascas. Memorias de Jeannel R. 1919b. Troglorites breuili, nouveau Carabique cavernicole des la Real Sociedad Española de Historia Natural, T. del cincuentenario Pyrénées espagnoles. Bulletin de la Société Entomologique de France, 509-539. 1918, 273-276.

548 New data and new subspecies of Troglorites breuili

Jeannel R. 1921a. Carabidae, p. 512-517 in: Bolívar y Pieltain C., Jeannel Ortuño V.M., Marcos J.M., Zabalegui I. 1996. Nuevos datos sobre R. Coleópteros cavernícolas nuevos de las provincias vascas. Memorias de Caraboidea de la Península Ibérica (3ª nota). Boletín de la Asociación la Real Sociedad Española de Historia Natural, T. del cincuentenario, española de Entomología 20: 31-39. 509-539. Ortuño V.M., Serrano J., Andújar A., Lencina J.L. 2003. Th e female Jeannel R. 1921b. Pselaphidae, p. 517-524 in: Bolívar y Pieltain C., genitalia of the genus Zabrus (Coleoptera: Carabidae: Zabrini). I. Th e Jeannel R. Coleópteros cavernícolas nuevos de las provincias vascas. general structure and the subgenera Zabrus, Euryzabrus, Platyzabrus Memorias de la Real Sociedad Española de Historia Natural, T. del and Epomidozabrus. European Journal of Entomology 100: 115-121. cincuentenario, 509-539. Rambla M. 1980. Arácnida, Opilionida, p. 529-533 in: Bellés X., Blas Jeannel R. 1942. Coléoptères Carabiques (2eme partie). Faune de France, M., Comas J., Escolá O., Español F., Estany J., Rambla R., Ribera 40. Lechevalier, Paris, 572–1173 p. C., Serra A., Vicente M. C., Vives E. Contribución al conocimiento de Jiménez-Valverde A., Ortuño, V.M. 2007. Th e history of endemic Iberian la fauna cavernícola del País Vasco. Kobie (Bilbao) 10: 525-568. ground beetle description (Insecta, Coleoptera, Carabidae): which Ribera C. 1980. Arácnida, Araneida, p. 534-538 in: Bellés X., Blas M., species were described ﬁ rst? Acta Oecologica 31: 13-31. Comas J., Escolá O., Español F., Estany J., Rambla R., Ribera C., Juberthie C., Decu V. 1994. Structure et diversite du domaine souterrain; Serra A., Vicente M. C., Vives E. Contribución al conocimiento de la particularites des habitats et adaptations des espèces, p. 5-22 in: fauna cavernícola del País Vasco. Kobie (Bilbao) 10: 525-568. Juberthie C., Decu V. (eds.), Encyclopaedia Biospéologica. Vol. 1. Salgado J. M., Blas M., Fresneda J. 2004. Nuevos datos sobre el género Société de Biospéologie, Moulis-Bucarest. Choleva Latreille, 1796 en la Península Ibérica con la descripción de Juberthie C., Delay B., Bouillon M. 1980a. Sur l’existence d’un milieu una nueva especie (Coleoptera: Cholevidae). Elytron [2003-2004] 17- souterrain superﬁ ciel en zone non calcaire. Comptes-rendus de l’Académie 18: 47-71. des Sciences de Paris 290(D): 49-52. Salgado J. M., Blas M., Fresneda J. 2008. Coleoptera, Cholevidae. In: Juberthie C., Delay B., Bouillon M. 1980b. Extension du milieu Ramos M. A., Alba J., Bellés X., Gosálvez J., Guerra A., Macpherson souterrain en zone non-calcaire: description d’un nouveau milieu E., Serrano J., Templado J. (Eds.). Fauna Ibérica, vol. 31. Museo et de son peuplement par les Coléopterès troglobies. Mémoires de Nacional de Ciencias Naturales. CSIC. Madrid. 802 p. Biospéologie 7: 19-52. Serra A. 1980. Chilopoda, Lithobiomorpha, p. 545-551 in: Bellés X., Blas Machado A. de B. 1940. A colecçao de aranhas cavernicolas do museu M., Comas J., Escolá O., Español F., Estany J., Rambla R., Ribera nacional de ciencias de Madrid. Las Ciencias (Anales de la asociación Contribución al conocimiento de la Española para el Progreso de las Ciencias) 7: 1-15. C., Serra A., Vicente M. C., Vives E. Magniez G. 1966. Contribution des spéléologues à la connaissance de fauna cavernícola del País Vasco. Kobie (Bilbao) 10: 525-568. certains Crustacés des grottes cantabres. Sous le Plancher 5: 30-41. Serrano J. 2003. Catálogo de los Carabidae (Coleoptera) de la Península Margalef R. 1953. Los crustáceos de las aguas continentales ibéricas. Instituto Ibérica. Monografías S.E.A.-9, Sociedad Entomológica Aragonesa, Forestal de Investigaciones y Experiencias, Ministerio de Agricultura, Zaragoza, 130 p. Madrid, 243 p. Vandel A. 1964. Biospéologie. La biologie des animaux cavernicoles. Gauthier- Mateu J. 1945. 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549 Appendix. Studied specimens and measures used in the morphometric analysis.

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).

Abbreviation 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. Measures in millimeters.

Nº TAXON SEX CAVE COLECTION HL HW MPW mPW PL HEW MEW EL

1 T. b. breuili ♂ A JF 1.58 2.02 2.4 1.54 2.06 2.06 3.27 5.67

2 T. b. breuili ♂ A JF 1.63 2.06 2.5 1.63 2.02 2.11 3.36 5.77

3 T. b. breuili ♂ A JF 1.68 2.06 2.5 1.68 2.06 2.16 3.36 5.76

4 T. b. breuili ♂ A JF 1.63 2.06 2.54 1.68 2.06 2.16 3.36 5.76

5 T. b. breuili ♀ A JF 1.58 1.92 2.3 1.53 1.82 1.92 3.07 5.19

6 T. b. breuili ♀ A JF 1.53 1.92 2.3 1.58 1.87 1.97 3.12 5.48

7 T. b. breuili ♀ A JF 1.49 1.97 2.45 1.58 1.87 2.06 3.22 5.52

8 T. b. breuili ♀ A JF 1.68 2.16 2.54 1.68 2.06 2.21 3.46 5.76

9 T. b. breuili ♀ A JF 1.68 2.11 2.5 1.68 2.02 2.11 3.36 5.76

10 T. b. breuili ♀ A JF 1.77 2.25 2.64 1.77 2.06 2.21 3.41 5.86

11 T. b. breuili ♂ M1 JF 1.49 1.82 2.26 1.53 1.92 2.02 3.17 5.48

12 T. b. breuili ♀ M2 JF 1.73 2.16 2.59 1.63 2.11 2.11 3.41 5.91

13 T. b. breuili ♀ A JF 1.68 2.11 2.5 1.68 2.06 2.11 3.31 5.62

14 T. b. breuili ♂ A JF 1.53 1.92 2.4 1.58 1.92 2.02 3.17 5.48

15 T. b. breuili ♂ A JF 1.73 2.06 2.59 1.73 2.11 2.16 3.46 5.91

16 T. b. breuili ♀ A JF 1.63 2.11 2.59 1.68 2.11 2.16 3.46 5.76

17 T. b. breuili ♂ A JF 1.73 2.21 2.64 1.73 2.11 2.3 3.41 5.96

18 T. b. mendizabali ♀ M JF 1.82 2.45 2.69 1.87 2.02 2.21 3.46 5.72

19 T. b. mendizabali ♂ Pa JF 1.97 2.59 2.78 1.87 2.26 2.5 3.75 6.15

20 T. b. mendizabali ♀ Pa JF 1.82 2.4 2.64 1.77 2.11 2.3 3.55 5.76

21 T. b. salgadoi ssp. n. ♂ V JF 2.21 2.78 3.27 2.3 2.16 2.59 3.89 6.34

22 T. b. salgadoi ssp. n. ♀ V JF 2.21 2.69 3.07 2.11 2.02 2.4 3.65 6.34 23 T. b. salgadoi ssp. n. ♀ V JF 2.16 2.69 3.17 2.11 2.06 2.35 3.84 6.05

24 T. b. salgadoi ssp. n. ♂ V VO 2.21 2.78 3.27 2.26 2.26 2.64 4.08 6.82 holot ype

25 T. b. salgadoi ssp. n. ♀ V VO 2.11 2.69 3.07 2.21 2.06 2.45 3.7 6.15

26 T. b. breuili ♀ A MNCNM 1.63 2.06 2.54 1.63 1.87 2.21 3.36 5.67 (Cot ype)

27 T. b. breuili ♀ A MNCNM 1.58 2.06 2.5 1.63 1.92 2.11 3.22 5.76

28 T. b. breuili ♀ A MNCNM 1.73 2.21 2.69 1.77 2.16 2.26 3.46 5.91

29 T. b. breuili ♂ M2 MNCNM 1.53 1.97 2.35 1.53 1.92 1.97 3.22 5.57

30 T. b. breuili ♂ M2 MNCNM 1.58 2.02 2.5 1.63 2.06 2.11 3.36 5.67

31 T. b. breuili ♀ M1 MNCNM 1.58 1.97 2.4 1.63 2.02 1.97 3.31 5.67

32 T. b. breuili ♀ M1 MNCNM 1.53 2.02 2.3 1.58 1.87 1.97 3.27 5.38

33 T. b. breuili ♀ B MNCNM 1.58 2.06 2.5 1.58 1.97 2.11 3.31 5.62

34 T. b. breuili ♂ A MNCNM 1.49 1.87 2.21 1.53 1.82 1.92 3.02 5.38

35 T. b. breuili ♀ A MNCNM 1.77 2.26 2.74 1.77 2.16 2.3 3.55 6.05

36 T. b. breuili ♂ A MNCNM 1.53 1.92 2.3 1.53 1.92 1.97 3.15 5.28

37 T. b. breuili ♂ A MNCNM 1.53 1.92 2.35 1.53 1.92 1.97 3.07 5.43

38 T. b. breuili ♂ A MNCNM 1.63 2.02 2.45 1.63 2.02 2.06 3.31 5.57

39 T. b. mendizabali ♂ Tx MT 1.82 2.26 2.64 1.87 2.06 2.21 3.41 5.67

40 T. b. mendizabali ♀ M MT 1.87 2.3 2.64 1.82 2.06 2.21 3.31 5.57

41 T. b. mendizabali ♀ M MT 1.87 2.3 2.64 1.82 2.02 2.16 3.36 5.72

42 T. b. mendizabali ♀ M MT 2.11 2.64 3.02 1.97 2.21 2.4 3.65 5.96

43 T. b. mendizabali ♀ M MT 1.97 2.5 2.88 1.97 2.11 2.26 3.5 5.96

44 T. b. mendizabali ♂ M MT 1.87 2.3 2.69 1.82 2.06 2.21 3.41 5.76

45 T. b. breuili ♂ A MT 1.53 1.92 2.4 1.68 2.02 2.02 3.22 5.52

46 T. b. breuili ♂ A MT 1.58 1.92 2.35 1.53 1.97 1.97 3.02 5.48

47 T. b. breuili ♀ Eb MT 1.63 2.06 2.5 1.63 2.02 2.11 3.31 5.52

48 T. b. breuili ♂ Eb MT 1.63 2.06 2.54 1.68 2.11 2.16 3.27 5.57

49 T. b. breuili ♂ Eb MT 1.77 2.21 2.69 1.73 2.3 2.35 3.46 5.91

50 T. b. breuili ♀ A MT 1.53 1.82 2.26 1.53 1.92 2.06 3.12 5.43

51 T. b. breuili ♀ A MT 1.63 2.06 2.54 1.63 2.06 2.16 3.36 5.76

52 T. b. breuili ♀ A MT 1.53 1.82 2.3 1.53 1.82 1.92 3.02 5.28 53 T. b. breuili ♀ A MT 1.58 1.92 2.45 1.58 1.97 2.02 3.17 5.43

54 T. b. breuili ♀ A MT 1.68 2.11 2.5 1.63 2.02 2.11 3.36 5.81

55 T. b. breuili ♂ M1 JS 1.58 1.97 2.45 1.68 1.97 2.06 3.27 5.48

56 T. b. breuili ♂ M1 JS 1.73 2.06 2.59 1.77 2.16 2.16 3.41 6

57 T. b. breuili ♂ M1 JS 1.73 2.16 2.64 1.73 2.11 2.21 3.41 6

58 T. b. breuili ♀ M1 JS 1.58 1.97 2.4 1.63 2.02 2.06 3.22 5.48

59 T. b. breuili ♀ M1 JS 1.68 2.06 2.45 1.63 1.97 2.16 3.41 5.67

60 T. b. breuili ♀ Et VO 1.68 2.02 2.45 1.63 1.97 2.11 3.27 5.62

61 T. b. breuili ♂ I VO 1.58 1.97 2.35 1.58 2.02 2.02 3.12 5.52

62 T. b. breuili ♀ I VO 1.58 1.92 2.35 1.49 1.92 1.87 3.12 5.28

63 T. b. breuili ♀ I VO 1.68 2.11 2.59 1.63 2.16 2.11 3.36 5.67

64 T. b. breuili ♀ I VO 1.73 2.11 2.54 1.63 2.06 2.11 3.36 5.67

65 T. b. breuili ♂ I VO 1.68 2.06 2.5 1.63 2.06 2.16 3.31 5.57

66 T. b. breuili ♀ I VO 1.73 2.11 2.5 1.77 2.06 2.16 3.31 5.57

67 T. b. breuili ♂ I VO 1.58 1.97 2.4 1.53 1.97 2.02 3.17 5.48

68 T. b. breuili ♀ I VO 1.63 2.02 2.4 1.63 1.97 2.02 3.12 5.43

69 T. b. breuili ♀ I VO 1.73 2.21 2.64 1.73 2.21 2.3 3.5 5.86

70 T. b. breuili ♂ I VO 1.63 2.02 2.45 1.58 2.06 2.16 3.22 5.62

71 T. b. breuili ♂ I VO 1.58 1.92 2.45 1.53 2.02 2.06 3.22 5.48

72 T. b. breuili ♀ I VO 1.73 2.11 2.54 1.63 2.06 2.16 3.31 5.67

73 T. b. mendizabali ♂ SZ VO 1.92 2.45 2.74 1.82 2.21 2.3 3.6 6.05

74 T. b. mendizabali ♀ Ek VO 1.92 2.5 2.88 1.92 2.26 2.5 3.7 6.15