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

), 2010, 46 (34) : 537-549

537
ARTICLE
New data on Troglorites breuili Jeannel 1919 (Coleoptera:
Carabidae: Pterostichini): a hypogean Iberian species
with description of a new subspecies
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.
Vicixri M. Oiruxo
(1)
, Javiii Fiisxiia
(2)
& Airuio 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
E-mail: vicente.ortuno@uah.es
Accept le 28 mai 2009
T
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
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).
538
V. M. Oiruxo, J. Fiisxiia & A. Baz
Tis 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 Ernio-
Zestoa 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. Te
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.
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, 15-07-
1998/26-04-1999, J. Fresneda leg. (V. M. Ortuo col.); 1,
idem, 18-07-2000, J. Fresneda leg. & col.
Genitalia study
Te 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.
Te study of the female genitalia involves a slightly more com-
plex procedure (see Carayon 1969; Ortuo et al. 2003): the
abdomen is dissected, and the contents are immersed in a satu-
rated KOH solution for 1218 hours. Membranous structures
are then stained with Chlorazol-E black. Te genital prepara-
tion 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. Te
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.
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).
A M1 M2 B Eb Et I M Pa Tx SZ Ek V
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
Table 2. Mean values and Standard deviations of the morphometric
variables used to discriminate subspecies.
T. b. breuili
n = 58
T. b. mendizabali
n = 11
T. b. salgadoi ssp. n.
n = 5
mean sd mean sd mean sd
HW 2.033 0.106 2.426 0.126 2.726 0.049
HEW 2.096 0.104 2.296 0.119 2.486 0.124
MEW 3.282 0.13 3.518 0.144 3.832 0.169
MPW 2.47 0.119 2.749 0.127 3.17 0.1
mPW _1 1.628 0.075 1.865 0.065 2.198 0.086
HL 1.627 0.079 1.905 0.087 2.18 0.044
EL 5.627 0.198 5.86 0.2 6.34 0.296
PL 2.014 0.1 2.125 0.092 2.112 0.097
New data and new subspecies of Troglorites breuili
539
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.
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).
Torax (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
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.
Figure 2
Head in ventral view of T. b. breuili from Cueva de Akelar (Navarra) (scale:
1 mm).
540
V. M. Oiruxo, J. Fiisxiia & 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 1
st
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 2
nd
stria,
from six to eight discal setae on the 3
rd
interstrial interval
and two or three on the 5
th
, one apical seta on 7
th
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. Te 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). Te 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).
New data and new subspecies of Troglorites breuili
541
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.
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).
Te 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. Tese
specimens are highly macrocephalous, accompanied
by a remarkable widening of the pronotum (Fig. 1c).
Te 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. Te 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. Te 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-
Table 3. Summary of the Discriminant Analysis.
Discriminant
function
Eigenvalue Percent
relative
Canonical
correlation
P-value
1 10.3436 91 0.955 0.000
2 1.022 9 0.711 0.000
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.
542
V. M. Oiruxo, J. Fiisxiia & A. Baz
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 di cult. Te 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. Te 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. Te derived classication functions (Table 4)
can be used to determine to which group each case
most likely belongs. Tere 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. Tat 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
Table 4. Coe cients of the classication functions for each one of the three
T. breuili subspecies.
T. b. breuili
T. b.
mendizabali
T. b. salgadoi
ssp. n.
HW 174.081 102.021 140.331
HEW 152.587 156.392 176.482
MEW 155.014 149.748 171.61
MPW 71.714 112.508 66.675
mPW _1 145.465 192.188 263.12
HL 167.941 207.298 262.152
EL 146.261 130.491 137.912
PL 34.715 56.006 159.825
CONSTANT 461.716 506.17 656.908
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.
New data and new subspecies of Troglorites breuili
543
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 Sagain-
Zelaia 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). Tis 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. Tis
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
Te 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. Tis a rmation
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. (Eurys-
peonomus) breuili (Jeannel 1919) [Coleoptera, Leiodi-
dae]. 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 rar-
est subspecies (collected as single individuals in each
exploration), and is observed exclusively under clay
heaps. In conclusion T. breuili is clearly a typical ele-
ment 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 hypo-
gean 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 prea-
dapted species Vandel 1964) but especially, by their
contribution of energy and materials (and their trans-
porters): it is an environment without solar radiation
and for this reason, without autotrophic species (Ju-
berthie & 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 speci-
mens (highly specialized troglobiomorph facies), are
very e cient in the exhaustive exploitation of the lim-
ited resources oered by the hypogean environment
(especially in deep areas in the caves). It should hap-
pen in the subspecies T. b. mendizabali and T. b. salga-
doi 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 occu-
pies the summit of the trophic pyramid? Its abundance
may suggest that, unlike the other two subspecies, it is
an opportunist organism, with higher adaptive capac-
ity, perhaps in agreement with its probable presence in
the more supercial hypogeous environment, the Mes-
ovoid Shallow Substratum (the MSS of Juberthie et al.
1980a, b). Tis 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 mov-
ing into the MSS. Te fact that this subspecies is able
544
V. M. Oiruxo, J. Fiisxiia & A. Baz
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
CAVES
REFERENCES M Er Tx SZ M2 Ek A Pu B Et Eb Al
Hirudinea
Herpobdellidae
Herpobdella octoculata (L. 1758) +
Galn (2006)
Gasteropoda
Zonitidae
Oxychillus arcasianus (Servain 1880) +
Galn (2006)
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)
+
+
+
+
+
+
+
+
Rambla (1980)
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.
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
Ribera (1980)
Galn (2006)
Machado (1940)
Isopoda
Oniscidae
Oniscus asellus L. 1758 +
Galn (2006)
Diplopoda
Glomeridae
Loboglomeris rugifer (Verhoe 1906) +
Galn (2006)
Chilopoda
Lithobiidae
Lithobius pilicornis doriae (Pocock 1890)
Lithobius validus vasconicus Chalande 1905
Lithobius tricuspis Meinert 1872
Lithobius romanus inopinatus Matic & Darabantzu
1968
+
+
+
+
Serra (1980)
Galn (2006)
New data and new subspecies of Troglorites breuili
545
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 re-
sources available, thermal and hygrometric conditions,
etc.). It is likely, but not certain, that the three subspe-
cies 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. Tis 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 di cult 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
TAXA
Troglophilic and regular trogloxenic organisms
CAVES
REFERENCES M Er Tx SZ M2 Ek A Pu B Et Eb Al
Diptera
Sciaridae
Lycosia sp.
Mycetophilidae
Tarnania fenestralis (Meigen 1818)
Culicidae
Culex pipiens L. 1758
+
+
+
Galn (2006)
Trichoptera
Limnephilidae
Micropterna nycterobia Mac Lachlan 1875 +
Galn (2006)
Lepidoptera
Geometridae
Triphosa dubitata (L. 1758) +
Galn (2006)
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
+
+
+
+
+ +
+ +
+
+ Vives (1980)
Espaol (1948)
546
V. M. Oiruxo, J. Fiisxiia & 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
CAVES
BIBLIOGRAFA M Er Tx SZ M2 Ek A Pu Al M1
Oligochaeta
Haplotaxidae
Haplotaxis navarrensis Delay 1973 +
Delay (1973)
Asellota
Stenasellidae
Stenasellus breuili (Racovitza 1924) +
Magniez (1966)
Copepoda
Ameiridae
Stygonitrocrella dubia (Chappuis 1937)
Nitocrella vasconica Chappuis 1937
+
+
Margalef (1953)
Isopoda - Oniscoidea
Trichoniscidae
Trichoniscoides pseudomixtus (Arcangeli 1934)
Trichoniscoides dubius (Arcangeli 1935)
Trichoniscoides cavernicola (Budde-Lund 1885)
+
+
+ +
+
+
Escol (1980)
Arcangeli (1935)
Galn (2006)
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.
+
+ + + +
+
+ +
+
+
+
Estany (1980)
Galn (2006)
Araneae
Linyphiidae
Troglohyphantes alluaudi Fage 1919
Micrargus cupidon (Simon 1913)
Leptyphantes bolivari Fage 1931
+
+ +
+
+ +
+
Ribera (1980)
Bells (1987)
Galn (2006)
Diplopoda
Vandeleumidae
Vandeleuma hispanica (Ceuca 1967)
Julidae
Mesoilus cavernarum (Verhoe 1938)
Mesoilus henroti (Mauris 1971)
Mesoilus stammeri (Verhoe 1936)
+
+
+
+
+
+
+
+
Ceuca (1967)
Vicente (1980)
Mauries (1971)
Galn (2006)
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)
New data and new subspecies of Troglorites breuili
547
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. Te 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. Te
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. Te
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.
Terefore 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). Tis
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
TAXA
Troglobitic organisms
CAVES
BIBLIOGRAFA M Er Tx SZ M2 Ek A Pu Al M1
Diplura
Campodeidae
Podocampa simonini Cond 1956
Litocampa espanoli Cond 1950
+ +
+
Cond (1956)
Galn (2006)
Coleoptera
Pselaphidae
Prionobythus bolivari Jeannel 1921 + +
Jeannel (1921b)
Coleoptera
Leiodidae
Numerous species
In the text
548
V. M. Oiruxo, J. Fiisxiia & 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.
Te 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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