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Revista Ibérica de Aracnología, nº 30 (30/06/2017): 3–10. ARTÍCULO
Grupo Ibérico de Aracnología (S.E.A.). ISSN: 1576 - 9518. http://www.sea-entomologia.org
A NEW LEIOBUNUM SPECIES FROM THE IBERIAN PENINSULA
(OPILIONES: SCLEROSOMATIDAE: LEIOBUNINAE)
Carlos E. Prieto1 & Hay Wijnhoven2
1 Departamento de Zoología y Biología Celular Animal, Universidad del País Vasco (UPV / EHU), PO box 644, 48080 Bilbao, Spain.
— carlos.prieto@ehu.eus
2 Groesbeeksedwarsweg 300, NL-6521 DW Nijmegen, Netherlands. — hayw@xs4all.nl
Abstract: A new Leiobunum species is described from Spain, Leiobunum incantaturbs sp.n. The species is characterised by its
colour pattern, large size and very long legs, the absence of denticle rows on the retrolateral sides of coxae I to III and its penial
morphology. It has been found in two separate areas, Serranía de Cuenca (Castilla-La Mancha) and Sierra de Gúdar (Teruel,
Aragón), both in the southern part of the Sistema Ibérico mountains. An isolated record is from the Spanish Pyrenees in Pobla de
Segur (province of Lleida, Catalonia).
Key words: Opiliones, Sclerosomatidae, Leiobuninae, Leiobunum, Cordillera Ibérica mountains, Pyrenees, Spain.
Una especie nueva de Leiobunum de la Península Ibérica (Opiliones: Sclerosomatidae: Leiobuninae)
Resumen: Se describe una especie nueva de Leiobunum de España, Leiobunum incantaturbs sp.n. Se caracteriza por su patrón
de coloración, gran tamaño y muy largas patas, la ausencia de hileras de dentículos en los lados retrolaterales de las coxas I-III y
por la morfología penial. Ha sido encontrada en dos áreas separadas, Serranía de Cuenca (Castilla-La Mancha) y Sierra de Gúdar
(Teruel, Aragón), ambas en el sur del Sistema Ibérico. Y se atribuye una localidad aislada de Pirineos, en Pobla de Segur (Lleida,
Cataluña).
Palabras clave: Opiliones, Sclerosomatidae, Leiobuninae, Leiobunum, Sistema Ibérico, Pirineos, España.
Taxonomía: Leiobunum incantaturbs Prieto & Wijnhoven n. sp.
Introduction
The subfamily Leiobuninae includes 13 genera of which three
are represented in Europe: Leiobunum C.L.Koch, 1839, Cos-
mobunus Lucas, 1848 and Nelima Roewer, 1910 (Kury,
2017). Although the genus Leiobunum has a Holarctic range
and includes well over one hundred species, recent analyses
(Hedin et al., 2012; Burns et al., 2012) have clearly demon-
strated that it is a polyphyletic genus, and this also applies to
Nelima, with several american and japanese Leiobunine and
Gagrelline genera as inner branches of the Leiobunum/Nelima
radiation. That leaves the taxonomy of the subfamily in a
chaotic state to be resolved in the future by the reassignment
of many ‘exotic’ species of Leiobunum or Nelima to their
proper, already created or new, American and Japanese gene-
ra. Fortunately, considering the European taxonomy the genus
name Leiobunum is the oldest and it was erected for a Euro-
pean species. Therefore, future nomenclatural changes for the
current European taxa are expected to be comparably few.
Until now seven Leiobunum species have been reported
from the Iberian Peninsula (Prieto, 2003; Prieto & Fernández
2007; Prieto 2008): L. argentipalpe Prieto & Fernández,
2007, L. blackwalli Meade, 1861, L. defectivum Rambla, 1959
(redescribed by Prieto & Fernández, 2007), L. granulosum
Prieto & Fernández, 2007, L. levantinum Prieto & Fernández,
2007, L. rotundum Latreillle, 1798 and L. lusitanicum
Roewer, 1923. Except for L. blackwalli and L. rotundum,
having large European distribution areas (Martens, 1978), the
remaining five species are considered endemics for the Iberi-
an Peninsula. In this contribution we present a new, most
likely endemic Leiobunum from Spain. It was first mentioned
and figured by Prieto (2008, Fig. 11) as Leiobunum sp. nov.
B. Several more representatives of Leiobuninae from the
Iberian Peninsula await a description (Prieto, 2008).
Material and Methods
Specimens were sampled by hand, immediately stored in
alcohol 70º and preserved in the collections indicated below.
Harvestmen bodies were photographed by combining
multiple images taken at different focus distances by a camera
attached to a Nikon SMZ1500 stereomicroscope and then
stacked with the Helicon Focus 6.2.2 software. Original
drawings were based on sketches directly drawn from an
Olympus stereo light microscope (100-1000x) with the aid of
a calibrated drawing mirror. All measurements are given in
mm. Coxal formula (Prieto & Fernández, 2007) is defined as
the presence of denticle rows on prolateral (front: F) or
retrolateral (back: B) sides of coxae I-IV, indicating weak
denticle rows with lower-case letters (f, b) and facultative
rows with apostrophe (f’, b’). BLI (Beinlänge Index [Staręga,
1972]) is the ratio between the length of femur I and the
cephalothorax width (at level of lobes between coxae II and
III).
Abbreviations
CHW: Collection of Hay Wijnhoven, Nijmegen, Netherlands.
CJM: Arachnological Collection of Jochen Martens, Mainz,
Germany.
ZUPV: Arachnological Collection of the Departamento de Zoología
y Biología Celular Animal, Universidad del País Vasco, Bilbao,
Spain.
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Taxonomy
Family Sclerosomatidae Simon, 1879
Subfamily Leiobuninae Banks, 1893
Genus Leiobunum C.L.Koch, 1839
Leiobunum incantaturbs new species
Leiobunum sp.nov. B – Prieto, 2008: 60, fig.11
TYPE LOCALITY. Cueva del Nacimiento near La Cueva,
municipality of Vega del Codorno (province of Cuenca,
Castilla-La Mancha), MGRS coordinates: 30T WK910765,
1400 m a.s.l. On walls and ceiling along the main passage
open at bottom of the large entrance of the cave (Fig. 1, 2).
TYPE MATERIAL.
Holotype. Male from type locality, C. Prieto, J. Fernández,
M. Higuera leg., 26.07.2004 (at walls of a cave). The speci-
men is deposited in the Museo Nacional de Ciencias Natura-
les, Madrid (MNCN 20.02/18019).
Paratypes. 1F [ZUPV 1922]; 6F, 15J [ZUPV 1920]; 3M, 8F,
11J [ZUPV 1921]: same data as the holotype. 1M, 5J [ZUPV
1917], La Ciudad Encantada (province of Cuenca, Castilla-La
Mancha), 30TWK842511, 1407 m a.s.l. (roof of a rock shel-
ter), C. Prieto, J. Fernández, M. Higuera leg., 25.07.2004.
2M, 1F [ZUPV 4705]; 2M [ZUPV 4706], Tragacete: Fuente
de San Blas (province of Cuenca), 30T WK993696, 1430 m
a.s.l., C. Prieto leg., 09.08. 2011 (roof of a road bridge). 1M,
1F [CHW; ex-ZUPV 4707]; 1F [ZUPV 4708], type locality,
C. Prieto leg., 09.08.2011 (walls and ceiling of the cave gal-
lery). 2M [ZUPV 5151] Cuevas de Almudén: Barranco de
San Pedro (province of Teruel, Aragón), 30T XL840088,
1260 m a.s.l., C. Prieto leg., 25.07.2014 (bridge under road).
2F [ZUPV 5152] Miravete de la Sierra: towards Aliaga (pro-
vince of Teruel, Aragón), 30T XK9501943, 1210 m a.s.l., C.
Prieto leg., 26.07.2014 (bridge under road).
ADDITIONAL MATERIAL. 2M [CJM 2674], La Pobla de
Segur: Congost de Collegats (province of Lleida, Cataluña),
31T CG38, 575 m a.s.l., W. Schawaller leg., 31.08.1980
(walls of an old tunnel).
DESCRIPTION:
Short diagnosis. Leiobunum incantaturbs n. sp. can be
differentiated from the remaining Iberian Leiobunum by the
following characters: retrolateral side of coxa I to III without
denticles, large body size (males 4.0-6.4) and very long legs,
body pale yellowish brown with dark spots including a black
saddle, ocularium light with narrow dark rings around the
eyes.
Male. Holotype. Body length 5.8, prosoma width 3.2, BLI
3.42. Leg lengths (Tab. I).
Prosoma. Yellowish with prominent dark brown to black
patch in front of the eye tubercle (Fig. 3). Prosoma margins
bordered with black. Supracheliceral lamellae extended into a
pair of blunt projections which bear a few indistinct black
denticles (Fig. 4D).
Dorsum. Ground colour of the dorsum pale yellowish with
contrasting dark brown to black pattern. Black saddle with a
pair of white spots or a row of up to eight spots per abdominal
segment. Dorsal integument with microgranulate microsculp-
ture.
Eye tubercle. Rather small (0,6 wide, 0,5 long), set at less
than twice its length from the frontal margin (Fig. 4B). About
as long as wide, slightly canaliculate, restricted basally and
slightly tilted backwards, smooth above with only some sen-
sory setae. Pale yellowish, eyes surrounded by black.
Venter and coxae. Pale yellowish, not contrasting with dor-
sum. Coxae smooth, with scattered sensilla chaetica. Rows of
denticles on prolateral side of coxae I to III and a row at the
retrolateral side of coxa IV; prolateral side of coxa IV with a
few indistinct denticles, resulting in coxal denticle formula F-
F- F- fB (Tab. II). Most denticles are rectangular with a tri-
dentate distal margin. Numbers of coxal denticles in the holo-
type (coxae I-IV, l/r): 16/14, -/-, 15/8, -/-, 24/23, -/-, 9/12,
19/19. Genital operculum with sensilla chaetica, near the top
with a few blunt denticles (Fig. 4C), microsculpture consist-
ing of pointed microgranulae.
Chelicerae. Smooth, pale yellowish, dorsal side of the second
segment provided with sensilla chaetica (Fig. 4A). Medially
first segment with an irregular row of sensilla chaetica, sec-
ond segment with extensive field of stout, short sensilla chae-
tica (Fig. 4A, left).
Pedipalps. Robust, pale yellowish (Fig. 5). On the ventral
region of the femur six tubercles occur, medially with an
additional indistinct row of four black-tipped denticles, base
with a short row of four denticles. The patella has black-
tipped denticles. Tibia long and slender, ventrally with few
small tubercles. Pedipalpal tarsus almost straight, very slightly
curved inwards near the top, ventrally with four small denti-
cles. Tibia and tarsus covered with trichomes. Claw pectinate
(Fig. 5C).
Legs. Long and slender (Tab. II; BLI= 3.42 in the holotype),
ground colour yellowish-brown, trochanters with dark-brown
patch on pro- and postlateral side, extending onto the proxi-
mal quarter of femora as longitudinal dark-brown stripes. Tips
of femora, patellae and tibiae whitish. Base of femora armed
in front and back region with a short row of denticles, most
prominent on leg femora I and II. Femora provided with ir-
regular rows of black tipped spines, accompanied by a sensil-
lum chaeticum.
Penis. Long and slender (Fig. 6). Base distinctly robust, wider
than alate portion. Truncus base tapering, then continuing
almost parallel-sided towards the alae. Intrinsic penial muscle
in about the proximal third of the penis. Proalate section short.
Alate portion well developed and elongated, almost parallel-
sided, consisting of two dorsal and two ventral membranes
which are fused in the proximal half section, forming two
pockets opening to the exterior distally as well as laterally.
Ventral membranes attached to the truncus far distally of the
dorsal ones, characteristically curved forming irregular folds
proximally. Dorsal membranes attached to the truncus more
proximally, 'intersecting' with the ventral membranes (from a
ventral or dorsal view) in a characteristic curve, extending on
to the ventral side and forming two small wings that partly
enclose the ventral membranes (Fig. 6A). Wings with internal
field of oval-shaped microgranulae. Glans long and tapering,
ventral margin concave (Fig. 6B), distally with two pairs of
sensory setae. Stylus robust and short, at its base with a dis-
tinct bump, tip terminating in a rounded bulge (Fig. 6G, H).
Measurements: penis length 2,60; basal truncus width 0,34;
truncus to pockets length 1,51; pockets to glans length 0,73;
pockets width 0,30; glans length 0,36.
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Fig. 1. Type locality of Leiobunum incantaturbs n. sp. Cueva del Nacimiento above La Cueva village, municipality of Vega del Co-
dorno (province of Cuenca, Castilla-La Mancha). Photo: vegadelcodorno.net // Fig. 1. Localidad típica de Leiobunum incantaturbs
n. sp. Cueva del Nacimiento, encima del pueblo de La Cueva, municipio de Vega del Codorno (Cuenca, Castilla-La Mancha). Foto:
vegadelcodorno.net
Fig. 2. Loose aggregation of Leiobunum incantaturbs n. sp. on the ceiling of the main passage from Cueva del Nacimiento (July 26,
2007). Note the abundance of small flies which probably were part of their diet. // Fig. 2. Agregación suelta de Leiobunum
incantaturbs n. sp. en el techo de la galería principal de la Cueva del Nacimiento (26/07/2007). Nótese la abundancia de pequeñas
moscas que probablemente formen parte de su dieta.
Table I. Leiobunum incantaturbs n. sp. Mean lengths and ranges of leg femora and legs of males (n=9) and females (n=11).
Legs I II III IV
Femur length
Males 10.9 (10.3-11.6) 17.6 (16.8-18.6) 10.8 (10.2-11.4) 13.6 (10.5-15.1)
Females 9.5 (8.8-10.3) 15.9 (14.9-17.8) 9.5 (9.0-10.4) 12.6 (11.9-14.0)
Total leg length
Males 48.7 (46-50) 84.6 (74-93) 52.0 (50-55) 69.6 (66-73)
Females 44.2 (41-53) 77.4 (72-85) 44.9 (40-49) 60.2 (54-65)
Table II. Leiobunum incantaturbs n. sp. Mean numbers and range of denticles on the prolateral (F) and retrolateral (B) side of the
leg coxae I to IV for males (n = 9) and females (n = 11).
Coxae I II III IV
Front/Back F B F B F B F B
Male 15.7 (11-20) − 13.2 (5-25) − 22.1 (12-28) − 10.8 (4-30) 18.2 (11-30)
Female 13.5 (9-19) − 8.6 (3-15) − 14.9 (3-20) − 3.7 (0-10) 16.3 (10-20)
Fig. 3. Leiobunum incantaturbs n. sp. A-B, male and female from Tragacete (ZUPV 4707). C, male from Cuevas de Almudén
(ZUPV 5151). D, female from Miravete de la Sierra (ZUPV 5152). / Fig. 3. Leiobunum incantaturbs n.sp.A-B,machoy
hembra de Tragacete (ZUPV 4707). C, macho de Cuevas de Almudén (ZUPV 5151). D, hembra de Miravete de la Sierra
(ZUPV 5152).
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Fig. 4. Leiobunum incantaturbs n. sp., male (drawings from specimens from the type locality). A, Right chelicera, left median, right
lateral view. B, Eye tubercle, left lateral view (right is front side), right dorsal view (top is front side). C, Genital operculum, left a detail
of denticles and microsculpture. D, Supracheliceral lamellae. Scale bars: A-D 0.5 mm, detail of C scale bar 50 μm.
Fig. 4. Leiobunum incantaturbs n. sp., macho (figuras de especímenes de la localidad típica). A, Quelícero derecho en vista medial
(izquierda) y lateral. B, Oculario, vista lateral (frente hacia la derecha) y dorsal (frente hacia arriba). C, Opérculo genital, con detalle
de los dentículos y la microescultura. D, Láminas supraquelicerales. Escalas: A-D, 0.5 mm; detalle de C, 50 μm.
Fig. 5. Leiobunum incantaturbs n. sp., male right pedipalp (specimen from the type locality). A, Lateral view. B, Median view. Scale
ba r 0.5 m m. C. T arsal claw. Scale bar 50 μm. / Fig. 5. Leiobunum incantaturbs n. sp., pedipalpo derecho del macho (espécimen de
la localidad típica). A, Vista lateral. B, Vista medial. Escala: 0.5 mm. C. Uña del tarso. Escala: 50 μm.
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Fig. 6. Leiobunum incantaturbs n. sp., penis (specimen from the type locality). A,Ventral view of distal region. B, Lateral view of
glans. C, Dorsal view of distal region. D, Ventral view. E, Lateral view. F, Dorsal view. G, Stylus tip, ventral view. H, Stylus tip, lateral
view. Scale bars: A-F 0.5 mm; G, H 50 μm. / Fig. 6. Leiobunum incantaturbs n. sp., pene (espécimen de la localidad típica). A, Vista
ventral de la región distal. B, Vista lateral del glande. C, Vista dorsal de la región distal. D, Vista ventral. E, Vista lateral. F, Vista
dorsal. G, Punta del estilo, vista ventral. H, Punta del estilo, vista lateral. Escalas: A-F 0.5 mm; G, H 50 μm.
Fig. 7. Leiobunum incantaturbs n. sp., female, seminal receptacles (specimen from the type locality). Scale bar 50 μm. / Fig. 7.
Leiobunum incantaturbs n. sp., hembra, receptáculos seminales (espécimen de la localidad típica). Escala: 50 μm.
Female (Fig. 3 B, D). Body length 4.5-8.2, prosoma width
2.8-3.3, opisthosoma width 3.7-4.8, BLI 3.0-3.3. Colouration
of the dorsum very similar as in the male. Females have
weaker denticle rows than males. The prolateral row of coxa I
and the postlateral row of coxa IV are distinct, while other
rows can be weak or absent, resulting in female coxal denticle
formula F- f- f- f'B.
Legs. Long and slender (Tab. II), armature of spines less
pronounced as in the male.
Seminal receptacles (Fig. 7). The seminal receptacles are
located in the sixth ovipositor segment.
ETYMOLOGY. The specific epithet 'incantaturbs' is a Latinized
composed noun (in nominative) based on one of the finding
places, La Ciudad Encantada, meaning The Enchanted City
(incantatum = enchanted; urbs = city, town). La Ciudad
Encantada is a famous geological site near the city of Cuenca,
consisting of characteristic karst formations, which date back
to the Cretaceous.
DISTRIBUTION (Fig. 8): The new species has been found in
the southern half of the so-called Iberian System (Sistema
Iberica), which is separated from the northern half by the
valleys of the rivers Jalón (Ebro tributary) and Henares (Tajo
tributary). The southern Iberian System comprises three main
sierras, Montes Universales / Serranía de Cuenca (1935 m and
1866 m) at the west and Sierra de Gúdar (2024 m) and Sierra
de Javalambre (2020 m) at the east. At present, Leiobunum
incantaturbs n. sp. has been found in the Serranía de Cuenca
9
Fig. 8. Leiobunum incantaturbs n. sp. geographical distribution. Note that the Pyrenean locality of L. incantaturbs is within the range
of L. rotundum (the dotted line shows its known southern distributional range, according to data summarized in Prieto & Fernández
2007). L. rotundum occupies also the northern part of the Iberian System, reaching the Monasterio de Piedra, just south of the Jalón
river. / Fig. 8. Leiobunum incantaturbs n. sp. Distribution geográfica. Nótese que la localidad pirenaica de L. incantaturbs está
dentro del área conocida de L. rotundum (la línea de puntos indica su límite meridional, según los datos recogidos en Prieto &
Fernández 2007). L. rotundum ocupa también la parte septentrional del Sistema Ibérico, alcanzando el Monasterio de Piedra, justo
al sur del río Jalón.
(three localities in the province of Cuenca, including the type
locality) and in the Sierra de Gúdar (two localities in the
province of Teruel), which lies a hundred km east from the
type locality. Two records from Cuenca (but not the type
locality) are located in the Parque Natural de la Serranía de
Cuenca.
Moreover, two specimens from a river canyon in the
northeastern part of the province of Lleida are also considered
conspecific, but not paratypical, because the Congost de
Collegats is more than 300km northeast of the type locality,
separated by the wide Ebro river basin; also there is a notable
altitudinal difference (1210-1430 m a.s.l. versus the Pyrenean
records at 575 m a.s.l.).
HABITAT: Most specimens of L. incantaturbs n. sp. are
collected from walls and ceilings of caves and under bridges
(Fig. 1, 2). It is expected they group together in these
sheltered spaces during day time to prevent desiccation and as
a protection against direct sunlight. Many other Iberian
Leiobunum species can be found at similar locations
(unpublished personal observations). In the type locality small
aggregations were recorded at the ceiling and the gallery
walls of the large cave and its entrance.
PHENOLOGY: Adults and subadults have been collected end
of July, adult specimens in the beginning of August,
suggesting that L. incantaturbs n. sp. matures in late summer.
It probably has a phenology common to many Central-
European and Mediterranean Leiobunum species, with
reproducing populations from late summer till late autumn or
early winter.
DISCUSSION: Leiobunum incantaturbs is clearly defined by its
penial morphology, in combination with the dorsal colour
pattern and the configuration of the coxal denticles rows.
Moreover, it is quite a large species, being averagely
somewhat larger than Leiobunum defectivum, the largest
Iberian Leiobunum so far, with legs I-IV reaching 50, 88, 53
and 69 mm (Prieto & Fernández, 2007). The absence of
denticles on coxae I to III suggests that it is not belonging to
the species-rich 'rotundum-group' having formula FB FB F-
FB with Iberian representatives: L. argentipalpe, L.
defectivum, L. lusitanicum, and L. rotundum and North-
African species L. biseriatum Roewer, 1910, L. coccineum
Simon, 1878 and L. cupreum Simon, 1878 (Prieto &
Fernández, 2007). Although till now few molecular data have
been published on European Leiobuninae (Hedin et al.,
2012), the provisional classification of the mentioned species
of the 'rotundum-group' as a natural group of closely related
species may turn out to be justified since there are also clear
similarities in penial morphology and some secondary
characters (postlateral denticle row coxa III lacking; male
mostly with uniform dorsal colouration, female more
contrasting; pedipalp, chelicera and genital operculum only
slightly sexually dimorphic).
The penial morphology of L. incantaturbs neither seems
to be related to the Iberian L. granulosum and L. levantinum
with which species it does share a similar coxal denticles
10
arrangement. In this respect the new species seems to stand
somewhat apart from other known Iberian taxa. From this it is
obvious that the European Leiobuninae should be thoroughly
revised based on molecular analyses. This is also clearly
demonstrated by a recent morphological revision of the four
species belonging to the Leiobunum rupestre species group
(Martens & Schönhofer, 2016).
The distribution of L. incantaturbs n. sp. can be charac-
terized as northeastern Iberian, occurring in the adjacent
provinces of Cuenca (Castilla-La Mancha) and Teruel
(Aragon). However, the harvestman fauna of the southern part
of the Iberian System is very poorly known: within the
distributional range of L. incantaturbs n. sp. in Cuenca no
other records of any Leiobunum species have been published
(Prieto & Fernández, 2007) and from the province of Teruel
no more than two records of Leiobunum blackwalli (Rambla,
1998) are documented.
In respect to the odd record in the Spanish Pyrenees
(Congost de Collegats, CJM 2647) far distant from the two
nuclei in the southern part of the Iberian System, we face a
similar paucity of records. There have been reported some
findings of Leiobunum 'biseriatum' from nearby localities
(Roewer 1925) which may refer to our new species. Possibly,
L. incantaturbs n. sp. is wide-spread in north-eastern moun-
tainous regions of Spain. Yet, because of the limited number
of records from the intermediate region, a basin with xeric
conditions between the Iberian and Pyrenean chains, this
remains to be confirmed.
Acknowledgements
The Basque Government through the Research group on
‘Systematics, Biogeography and Population Dynamics’ (IT575-13)
and the Spanish Ministerio de Ciencia e Innovación (Ref.CGL2008-
01131/BOS) have funded the sampling for this work. HW expresses
his sincere gratitude towards the Uyttenboogaart-Eliasen Stichting
for awarding him with a grant (SUB.2012.05.04) to visit CP at the
Basque University in Bilbao as part of a collecting expedition to
Morocco and Spain. Jochen Martens kindly provided specimens
from his collection.
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