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A new orophilous species of the genus Dasylobus (Opiliones: Phalangiidae) from Sierra Nevada, Spain
Rocı
´o Martı
´nand Carlos E. Prieto
1
: Laboratorio de Aracnologı
´a y Malacologı
´a, Departamento de Zoologı
´a y Biologı
´a
Celular, Universidad del Paı
´s Vasco, Apdo. 644 48080-Bilbao, Spain
Abstract. A new species, Dasylobus nevadensis, is described; it inhabits high areas of the Sierra Nevada, the highest
mountain chain in Spanish mainland. The new species is smaller than all other Iberian Dasylobus species and has short,
annulated legs, light silver coloration, and is juvenile-like. Together with Roeweritta carpentieri (Roewer 1953), this is an
orophilous endemic species from Sierra Nevada.
Keywords: Iberian Peninsula, Phalangiinae, new species, taxonomy, Europe
The subfamily Phalangiinae comprises around 25 genera
within the Holarctic region, with high representation in
tropical Africa (Martens 1978; Stare˛ga 1984; Crawford
1992). In the western Mediterranean area, the subfamily is
represented by only three genera (Stare˛ga 1984), Phalangium
Linnaeus 1758, Metaphalangium Roewer 1911, and Dasylobus
Simon 1878, as well as four endemic Macaronesian genera,
Bunochelis Roewer 1923, Metadasylobus Roewer 1911, and
Parascleropilio Rambla 1975 from the Canary Islands, and
Ramblinus Stare˛ga 1984 from Madeira Island.
The genus Dasylobus is characterized by a unidentate
supracheliceral lamina, a basichelicerite with a dorsal granulated
mound, a palpal patella with a mesal, conical/rounded, hairy
apophysis, medium to long legs with an incrassate leg I, and a
penis with a broad basis, slender trunk and wedge-shaped glans,
and sometimes with dorsodistal spoon (Stare˛ga 1976; Martens
1978). Additionally, it has a denticulate carapace, frontal corners
and, sometimes, a saddle with a median lighter broad band.
In the Iberian Peninsula the genus is represented by several
species, although the status of some of them is not very clear.
Dasylobus echinifrons Simon 1879 was described from
Narbonne, La Clape and Le Vernet (France) although Simon
(1879) also reported it from Aranjuez and Sierra Morena
(Spain). The known range of D. graniferus (Canestrini 1871),
extends from Yugoslavia, Austria, Switzerland, Italy and
France to Spain (Chemini 1989), where it was reported as
Eudasylobus nicaeensis (Thorell 1876) from Toledo (Roewer
1923), a synonym of D. graniferus according to Chemini
(1986), and several sites from Sierra de Guadarrama (Simon
1879; Rambla 1967; Martens 1978). Dentizacheus ibericus
Rambla 1968 was described from Torre de Moncorvo
(Portugal) and transferred to Dasylobus by Prieto (2003).
Three other nominal species have been removed from
Dasylobus:D. lusitanicus Roewer 1956, which was described
from Coimbra (Portugal) but Stare˛ga (2004) redescribed it as
Metaphalangium lusitanicum, and Eudasylobus rondaensis
Kraus 1959, described from Sierra del Oreganal near Ronda
(Ma´laga) and localities in Tarragona (Sierra de Monsech) and
Murcia (Sierra Espun˜a), which was transferred to Dasylobus
by Prieto (2003) and synonymized with M. lusitanicum by
Stare˛ga (2004). The third nominal species, De. zuluetai
Rambla 1959, described from El Escorial (Madrid), was
synonymized with Da. echinifrons by Stare˛ga (1973) who later
(Stare˛ga 2004) placed it in the synonymy of M. lusitanicum.
The Sierra Nevada (Fig. 1), located in Andalusia between
Granada and Almerı
´a provinces, is the highest mountain
range on the Iberian Peninsula, reaching 3,478 m in the
Mulhacen and 3,395 m in the Veleta. The chain was formed
during the Alpine Orogeny, following the collision between
African and European plates, and the central massif is mainly
composed of heavily deformed metamorphic rocks, mainly
mica schists, locally with gneiss, quartzite and amphibolite
(Lo´pez-Bermu´dez et al. 1989). The highest part of the range is
included in the Sierra Nevada National Park.
Opilionological knowledge about the Sierra Nevada is very
scarce because only two species have been previously recorded:
Homalenotus coriaceus (Simon 1879) recorded by Kraus (1961)
from Rı
´o Monachil (30SVG60, 2,300 m) and Roeweritta
carpentieri (Roewer 1953), an endemic monotypic genus
recorded from six localities between 2,000 and 3,000 m
(Rambla 1960; Marcellino 1967; Barea 2008). During a trip
to the region in November 1982, we found many small
harvestmen under schistose stones. We routinely considered
these specimens as juveniles during sorting, but on close
inspection determined that they are adults belonging to an
undescribed species, which is described here.
METHODS
Taxonomic methods follow outlines proposed by Pinto-da-
Rocha et al. (2007). Body (carapace) width was measured
between the incisions of coxae II and III. BLI index is the
relation of the femur I length to the carapace width. All
measurements are in mm.
Specimens were studied, photographed and drawn with a
Nikon SMZ-1500 stereomicroscope provided with a drawing
tube and a digital camera. The penis and spermathecae were
drawn with a Nikon Optiphot. Photo stacks were combined
with the software Helicon Focus, and backgrounds were
cleaned with Photoshop.
The specimens studied in this contribution are lodged in the
Museo Nacional de Ciencias Naturales, Madrid (MNCN)
(holotype and female paratype) and the Departmento de
Zoologı
´a y Biologı
´a Celular, Universidad del Paı
´s Vasco,
Bilbao (ZUPV) (remaining paratypes).
TAXONOMY
Family Phalangiidae Latreille 1802
Genus Dasylobus Simon 1878
Dasylobus Simon 1878:ccxviii (footnote).
1
Corresponding author. E-mail: carlos_prieto@ehu.es
2010. The Journal of Arachnology 38:113–118
113
Eudasylobus Roewer 1911:53. Synonymized by Chemini
(1989).
Euplatybunus Roewer 1912:252. Synonymized with Eudasylo-
bus by Stare˛ ga (1984).
Parazacheus Lerma 1952:7. Implicitely synonymized with
Dasylobus by Chemini (1989).
Type species.—Opilio argentatus Canestrini 1871 by original
designation.
Diagnosis.—Penis shaft continuously narrowed from the
basal to the middle part, thence distad almost parallel or little
divergent edges, and moderately compressed dorsoventrally.
Glans strongly compressed, swollen underneath and plain
above. Palp femur smooth, patella with a mesodistal, short,
blunt conical apophysis, tibia and tarsus smooth and hairy.
Ocularium small, separated by half of its length from the
frontal border. Supracheliceral laminae with a pointed granule
on each (Martens 1978:290–291 for Eudasylobus).
Remarks.—Chemini (1989) reviewed the Italian taxa,
concluding that only four species occur in Italy, but the
remaining species are poorly known. According to Hallan &
Kury (2009), the genus comprises about 20 species distributed
within the Mediterranean region, from Asia Minor and
Lebanon to Algeria and the Iberian Peninsula, including the
larger islands (Cyprus, Crete, Sardinia, Corsica, Balearic), but
some entries are doubtful or have been removed recently to
synonymy. The Spanish fauna is currently composed of four
taxa (Stare˛ ga 2004; Prieto 2008); D. echinifrons (Simon 1879),
D. graniferus (Canestrini 1871) and D. ibericus (Rambla 1968)
from the Spanish mainland, and D. ferrugineus (Thorell 1876)
from Balearic Islands.
Dasylobus nevadensis new species
(Figs. 2–12)
Type material.—SPAIN: Granada: holotype male, Sierra
Nevada: road to the Veleta peak (UTM [WGS84]:
30SVG658050), 2,560 m, under stones in a stony slope, 2
November 1982, C.E. Prieto and A. Prieto (MNCN 20.02/
17103). Paratypes: 9 males, 11 females, 1 juvenile, same data as
holotype (ZUPV/0258bis); 1 female, same data as holotype
(MNCN 20.02/17104).
Figure 1.—Known localities of Iberian Dasylobus species and Metaphalangium lusitanicum.E,D. echinifrons (*, locality of the synonym
Dentizacheus zuluetai); G, D. graniferus;I,D. ibericus;N,D. nevadensis new species; L, M. lusitanicum (*, localities of the synonym Eudasylobus
rondaensis). Right bottom inset, western part of Sierra Nevada National Park with the type locality of D. nevadensis new species (white dot).
114 THE JOURNAL OF ARACHNOLOGY
Figures 2–12.—Dasylobus nevadensis new species: 2. Male holotype, dorsal view; 3. Female, dorsal view; 4. Male palp, internal view; 5. Male
palp tibia and patella, dorsal view; 6. Female palp, internal view; 7. Female palp tibia and patella, dorsal view; 8. Male chelicerae, lateral view; 9.
Penis, lateral view; 10. Glans, lateral view; 11. Glans, frontal view; 12. Spermathecae in female ovipositor. Scale bars 51 mm (Figs. 2–4, 6, 8),
0.5 mm (Fig. 9), 0.1 mm (Figs. 5, 7, 10–12). Figs. 3–12 from male and female paratypes.
MARTI
´N & PRIETO—A NEW OROPHILOUS DASYLOBUS FROM SPAIN 115
Etymology.—The specific epithet refers to the Sierra
Nevada, the highest mountain chain on the Iberian Peninsula,
which this species inhabits.
Diagnosis.—Belonging to the genus Dasylobus, this species
can be recognized by its juvenile appearance, small size (3.0–
3.9 mm in males and 3.5–4.5 mm in females), short (BLI
index, 0.87–1.01 in males and 0.69–0.81 in females), clearly
annulated legs, and palpal patella with a conical apophysis.
Description.—Male holotype: body length 3.76 mm, cara-
pace width 2.26 mm, femur I 2.04 mm, BLI 0.90.
Carapace (Fig. 2):dorsum smooth except for some scat-
tered denticles, mainly concentrated between ocularium and
ozopores. Supracheliceral laminae with one denticle on each
lamina. Frontal edge regularly concave, completely unarmed
in the center and with tufts of strong denticles on each anterior
corner. Ozopores visible from above and with 1–3 denticles
near the anterior and posterior corners. Ocularium silver in
color, except for dark ocular rings; medium-sized (1/4 of width
and 1/3 of length of carapace) with a medial groove and a row
of 7–8 denticles in each side around the eyes. Carapace divided
by a ledge, parallel to lateral borders, in a central area and two
lateral ones; lateral areas with a marginal silvery stripe and
four brown patches; central area with two brown close parallel
stripes on the preocular region, lateral sides with brown
patches separated by silvery color and two triangular patches
behind the ocularium. First and second thoracic tergites each
with a row of small denticles.
Abdomen: cream ground color. Abdominal scutum smooth,
but with several brown sclerotization spots. Dorsal saddle
wide, brownish but spotted in white, extending from first
thoracic tergite to area V, narrowed in areas I and IV, with
area I lined with silvery stripes. Remainder of abdominal
surface with brownish dots and whitish halos. Silvery anal
operculum. Ventral side cream-colored. Leg coxae smooth and
covered with numerous short setae, and fields of brown spots
on posterior faces. Genital operculum with irregularly
arranged short setae and 2 pairs of close brown spots basally.
Posterior border of genital operculum indicated by a brownish
‘M’-shaped patch. Abdominal sternites smooth, with scattered
setae and rows of brown patches on anterior edges.
Chelicerae (Fig. 8):first segment short, thickened, and with
a large dorsal protuberance with irregular surface, more
elevated on internal side and covered by granules and
scattered setae. Second segment with a dorsal domed
protuberance with frontal side covered by granules and setae.
Palps (Figs. 4, 5):trochanter with some dorsal and ventral
granules distally. Femur with a ventral field of setae and scarce
granules, a basal mesodorsal granular field, a dorsal row of
setae and few denticles, and a small distomesal thickening
covered by setae. Patella with a distomesal conical apophysis,
mesal side densely covered by a setose field, which continues
to the apophysis. Tibia with a very small setose distomesal
swelling. Tarsus densely covered by setae, mesal side with two
inconspicuous rows of microgranules. Femur to tibia each
with a dorsal light brown stripe.
Legs: relatively short (BLI 50.90). Femur, patella, and
tibia of leg I thickened. Femur with a whitish basal ring
(suture line for autotomy), rounded in transversal section and
with five rows of setose granules (13–17 in femur I). Patella
with rows of setae and 3 denticles on distodorsal edge. Tibia
with 3 dorsal rows of setae and 2 ventral rows of sharp
granules, and a small basodorsal denticle; spiracle located
retrobasally. Metatarsus with some granules on ventral side
and a pseudoarticulation at the distal third. Tarsus with many
tarsomeres (26, 45, 28–29, 31–32 in legs I–IV, respectively).
Legs brownish, with white annulations located as follows:
femur with basal, central and apical rings; apical on patella;
central and apical on tibia; basal and surrounding the
pseudoarticulation on tarsus.
Penis (Figs. 9–11):1.75 mm long. Shaft widened basally
(0.3 mm), strongly tapering until half of its length (0.1 mm
wide), then very gradually widened towards distal end
(0.15 mm wide). End of shaft without dorsal excavation.
Glans with triangular profile, and apically dilated in frontal
view; stylus with a secondary internal point.
Female (Figs. 3, 6, 7):similar to male but abdomen wider
and longer, and saddle less profiled. Chelicera normally
developed. Palpal patella (Fig. 7) with a more developed
apophysis and mesal side covered by longer setae; distomesal
apophysis of the tibia more developed and covered by longer
setae.
Spermathecae (Fig. 12):long and slender, bent toward the
ovipositor axis and extended until the third complete ring,
with a small secondary pouch on the first third of its length.
Measurements: see Tables 1 and 2.
Remarks.—The new species has male chelicerae with a large
dorsal protuberance on the basal article, palp with a
conspicuous distomesal apophysis on the patella and another
apophysis on the tibia (much smaller in the male), femur I
thickened, anterior corners of the carapace with tufts of strong
denticles, and supracheliceral laminae with a denticle on each.
These features indicate that the species belongs to the genus
Dasylobus.
Dasylobus nevadensis is easily distinguished from the other
Iberian species. Dasylobus ibericus and D. echinifrons are
bigger (body size 6 mm or larger) and robust, with longer and
Table 1.—Leg measurements and tarsal counts of the male holotype and a female paratype of Dasylobus nevadensis.
Trochanter Femur Patella Tibia Metatarsus Tarsus Total leg No.tarsomeres
Male Leg I 0.32 2.04 0.78 1.90 2.20 3.80 11.04 26
Leg II 0.38 3.57 1.00 3.15 3.29 7.03 18.42 45
Leg III 0.27 2.13 0.80 1.82 2.63 4.16 11.81 29
Leg IV 0.37 2.92 0.83 2.38 3.68 5.52 15.70 32
Female Leg I 0.29 1.65 0.70 1.50 1.75 3.01 8.9 27
Leg II 0.37 3.14 0.92 2.77 2.85 5.86 15.91 43
Leg III 0.29 1.79 0.66 1.55 2.05 3.36 9.70 27
Leg IV 0.32 2.61 0.74 2.05 3.05 4.39 13.16 32
116 THE JOURNAL OF ARACHNOLOGY
non-annulated legs (Simon 1879; Rambla 1968). Dasylobus
graniferus is distinguished by the presence of a conspicuous
protuberance above the articulation of cheliceral fingers on
the second cheliceral segment of the male (Rambla 1967;
Martens 1978; Chemini 1989).
Metaphalangium lusitanicum, formerly in Dasylobus, has
some features in common with Dasylobus, but has a short and
wide distal apophysis on the palpal patella, longer legs,
denticle rows on the ocularium with only 4–5 denticles, basal
segment of male chelicerae without dorsal denticulate bump,
and lacks a distal apophysis on the palpal tibia.
Another piece of data that partly justifies the specific
separation is the date of collection of the specimens; adults of
Metaphalangium lusitanicum were collected in March and
April (Kraus 1959, as Eudasylobus rondaensis), while the
specimens of Dasylobus nevadensis were gathered in Novem-
ber, which implies different life cycles for both species.
Curtis & Machado (2007) review the temporal patterns in
harvestmen, and Rambla (1985) and Tsurusaki (2003) discuss
phenological patterns for several northern or montane species.
Most montane and alpine species show hatching and growth
in late spring/summer, and maturation and egg-laying in late
summer/autumn, and finally hibernating over winter in the egg
stage. Dasylobus nevadensis may match that life cycle; in fact,
it matches the cycle of Harmanda nigrolineata Martens 1987,
which occurs in the Himalayan Mountains between 2,400 and
3,500 m (Curtis & Machado 2007, based on data from
Martens 1984). According to an ombrotermic diagram for
an Astronomic Observatory from the Sierra Nevada at
2,507 m and less than a kilometer from the type locality
(Rivas-Martı
´nez et al. 1997), there is a long period between
November and April when the temperature is below 0 uC and
the terrain is usually fully covered with snow that has fallen
during autumn and winter. Therefore, as we stated earlier, this
species would hatch, grow, and mature between May and
October–November.
On the contrary, only a few harvestmen with distributions
throughout southern/xeric regions have been studied. Tra-
chyrhinus marmoratus Banks 1894 from the arid regions of
Texas is a good example, with the adult stage between January
and July, with abundance peak in April (Curtis & Machado
2007). Another more relevant example is D. graniferus from
southern Italy (Chemini 1989); all data for localities in
Calabria and Sicily lie between May and early July.
Metaphalangium lusitanicum matches this southern life cycle,
with hatching and growth occurring in winter, and maturation
and egg-laying in spring, due to the strong and longer xeric
season of summer/autumn, as the phenological profile of
Opilio insulae Roewer 1956 in lower elevations in Crete
(Chatzaki et al. 2009), a species with a high phenological
plasticity because it lives also at 2,000 m, where the snow
cover lasts generally from November to May, matching the
phenological profile of Dasylobus nevadensis.
ACKOWLEDGMENTS
We are very grateful to Dr Mark Harvey, subject editor of
the Journal of Arachnology, and two anonymous reviewers
for their comments and suggestions that improved the final
version of this paper.
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Males (n59)
Average 3.49 2.04 1.89 10.18 17.48 11.16 14.61 0.92
Minimum 3.06 1.85 1.62 8.73 14.43 9.24 12.47 0.87
Maximum 3.90 2.26 2.09 11.19 18.55 12.68 15.7 1.01
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MARTI
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Manuscript received 14 January 2009, revised 2 November 2009.
118 THE JOURNAL OF ARACHNOLOGY
