Revision of the genus Dicranopalpus from northern Spain and Corsica, with descriptions of two new species (Arachnida, Opiliones, Phalangioidea)

. The Iberian Peninsula represents a diversity hotspot for the genus Dicranopalpus . However, most taxa are insuf ﬁ ciently de ﬁ ned. Our revision of the Dicranopalpus pyrenaeus species group reveals two hidden species: D. catariegensis sp. nov. and D. gallaecicus sp. nov. Two species, D. pyrenaeus from the Spanish and French Pyrenees and D. insignipalpis from Corsica, are redescribed. Dicranopalpus cantabricus Dresco, 1953 is resurrected as a valid species. Three new synonyms of D. martini (Simon, 1878) are proposed: Fagea bolivari Dresco, 1949 (including D. bolivari sensu Rambla 1975), Egaenasser extraordinarius Roewer, 1953 and Dicranopalpus dispar Rambla, 1967. With these additions, the Iberian Peninsula harbours four species of the Dicranopalpus pyrenaeus species group. They largely have allopatric distributions, ranging from the Pyrenees and the Cantabrian mountain range to the west coast of Galicia.

All species in the genus Dicranopalpus have sexually dimorphic pedipalps with a knob-shaped apophysis proximally on the femur and a long to extremely long apophysis on the patella medially, giving the pedipalp a forked (from the Greek 'dicranos', meaning 'fork') appearance (e.g., Martens 1978;Wijnhoven & Prieto 2015). Several species have an additional smaller apophysis medially on the tibia (e.g., D. gasteinensis Doleschall, 1852;Martens 1978). The patellar apophysis is smaller and more slender in males than in females. On their function, Wolff et al. (2016: 37) remarked: 'Apophyses may increase the area covered by glandular setae and may form a capture basket together with the fl exed tibia and tarsus. They are a barrier for a prey held between the pedipalps. This may help in restraining and holding down struggling prey.' In females, the pedipalp femur, patella (including the apophysis) and tibia have a dense cover of plumose setae, while males have sensilla chaetica instead (Wijnhoven 2013;. The areas with plumose setae are slightly infl ated to contain glandular tissue. The Iberian Peninsula (including the Spanish-French Pyrenees), with six valid species of Dicranopalpus , is particularly diverse, fi ve of which may be considered endemic to the peninsula and adjacent regions: D. bolivari (Dresco, 1949), D. caudatus Dresco, 1948, D. martini (Simon, 1878), D. pulchellus Rambla, 1960 andD. pyrenaeus Dresco, 1948. In the last decennia, D. ramosus has colonised northern parts of Spain and expanded to western and northern Europe (Wijnhoven & Prieto 2015). Dicranopalpus caudatus and D. pyrenaeus have been recorded from French Pyrenean regions as well (e.g., D'Amico 1988;D'Amico & Besson 1995;Delfosse & Melotti 2016;Delfosse 2017).
Within the group of species, which we here refer to as the Iberian assemblage, two 'species groups' can be distinguished, mainly based on pedipalp and penis morphology. The fi rst group consists of D. ramosus and D. caudatus , which have a globose glans and a short stylus with a brush of minute setae at its base (Martens 1978;Wijnhoven & Prieto 2015). The remaining four species have a different glans structure, a pair of horn-shaped projections on the dorsal side near the glans-stylus junction and a long, curved and sclerotized stylus. The second group comprises D. bolivari and D. pulchellus, displaying extreme sexual dimorphism, with rather 'normal' females, while males are distinctive for their pedipalps with large tubercles, long and slender patellar apophysis and robust chelicerae. A third group involves D. martini and D. pyrenaeus , referred to as the Dicranopalpus pyrenaeus species group or Dicranopalpus pyrenaeus group. The male pedipalps have a short patellar apophysis, approximately as long as the patella, and this feature also applies to D. insignipalpis (Simon, 1879) , endemic to Corsica. Two Italian species, D. larvatus (Canestrini, 1874) and D. brevipes Marcellino, 1970, reviewed previously (Wijnhoven & Martens 2019Wijnhoven et al. 2020), also show affi liation to this group. Taxonomy within the Dicranopalpus pyrenaeus group is complex, which can be illustrated by analysing the historical accounts on these species: D. martini , D. insignipalpis and D. pyrenaeus .
du second au quatrième segment, un peu atténuée en avant, élargie transversalement sur le troisième segment''; Simon 1878). Next, Roewer (1923) included D. martini in his world catalogue, commenting that he had not seen the type specimen. Moreover, Mello-Leitão (1936), without providing any further information, quoted the species for the fi rst time for Spain, in Vall d'Aran and Llívia (Cataluña). Dresco (1948) revised for the fi rst time the then known species of Dicranopalpus by studying the types of Simon's collection housed in the Paris MNHN, and new material and including an identifi cation key, mainly based on the shape of the pedipalps. About the holotype of D. martini Dresco (ibid.) wrote: ''The specimen in the Simon collection, tube 2493, has its venter opened and the ovipositor is lost. The animal is adult but its sex cannot be verifi ed. The tube has no identifi cation label, but Simon's catalogue mentions for that number: Prosalpia martini Sim., Portalegre (Martin). An additional juvenile specimen is found in the tube'' (''Le spécimen de la coll. Simon, tube 2493, a eu le ventre ouvert et l'ovopositor a disparu; l'animal est adulte mais le sexe ne peut etre vérifi é. Ce tube est sans étiquette de détermination, mais le catalogue Simon porte à ce numéro: Prosalpia martini Sim., Portalegre (Martin). Une bête non adulte se trouvait également dans ce tube''). In his key, he wrote: ''Femoral apophysis one third as long as the diameter of the femur'' ('apophyse basale infère du fémur de la p. m. atteignant le tiers du diamètre de l'article'; Dresco 1948), and he illustrated this character (Dresco 1948: 340, fi g. 7), this being the only published illustration of the holotype.
The next published records are from Roewer (1953), referring to two non-adult females from 'Monte de los Sagredales' (Asturias). He did not know the Mello-Leitão (1936) publication and stated that it was the fi rst record for Spain.
In summary, several nominal species are involved, of which none has been appropriately defi ned. In this paper, the results of a revision on D. insignipalpis and species of the Dicranopalpus pyrenaeus group are presented, based on older and newly collected unpublished material. Signifi cant characters for species identifi cation include the morphology of the penis and pedipalp. In total, two species are redescribed, two new species are identifi ed and described, and one species is resurrected. A key to the males of the Dicranopalpus pyrenaeus group is provided.

Material and methods
We collected the specimens by hand (mostly by turning stones and pieces of dead wood), stored them in 70% ethanol and preserved them in the collections mentioned below. Additional samples were provided from some other collections. Despite continuous efforts for years, we have not been given permission to investigate the material of Dicranopalpus from the Dresco and Simon collections of the Muséum national d'histoire naturelle in Paris.
For examination, the dissected penis was placed on a concave cavity slide and immersed in glycerine. It was covered with a cover glass that could be moved, facilitating the rotation of the penis while under the microscope. For each species a lateral, dorsal and ventral view of the glans is presented. For seminal receptacles, the ovipositor was cleared for some hours in a 5% KOH solution at room temperature. Original drawings were based on sketches directly drawn from an Olympus stereo light microscope and a Euromex dissecting microscope, with the aid of a calibrated drawing mirror.
Body length is measured from the anterior prosoma margin to the posterior margin of the opisthosoma. The BLI (body-leg index) is defi ned as length femur I/width prosoma at the level of leg coxae II and III (Staręga 1972). For names and numbering of muscles, and proposed muscle functions Shultz (2000) is followed. Defi nitions of sensory structures are according to Wijnhoven (2013) and Willemart et al. (2009). Measurements are given in mm. Penial morphology: the penis consists of a truncus, glans and stylus. The truncus base is a dark brown, sclerotized, semi-circular structure, with a lateral extension, curving ventrally and apically on each side. At the extensions, extrinsic penial muscles attach (Fig. 3B), which act in protracting, guiding and retracting the penis. It is of notice that during preparation for taxonomic purposes, these lateral extensions often break off, or even the complete base of the truncus gets torn off (Fig. 8B). The truncus is sclerotized, long and tubular, its proximal portion being wider and containing the intrinsic penial muscle (Fig. 3A). A thin seminal duct enters the truncus from the base (Fig. 3B) Fig. 3C-D). The large tendon of the intrinsic penial muscle is attached to the ventral glans base (Fig. 3D).

Abbreviations
The glans is globular to elongated, movable to some extent, and fl exed against the truncus. Glans dorsally with two sclerotized ridges widely spaced distally, forming a cavity with infl atable bladder ( Fig. 3C-D). Thick-walled, sclerotized areas are recognisable by interspersed canaliculi. The S-curved stylus has a pair of horn-shaped, forward-directed projections near the base. In two species ( D. insignipalpis and D. pyrenaeus ) these horns are fl attened and triangular (Figs 8C-I, 20E-K). Minute ducts of an unknown nature run from the glans to the horn tips. Laterally in the distal half of the glans there are two pairs of sensory setae. Presumably, these are present in all species under study, but because of their minute size they could not always be recognised (e.g., Figs 8F, H-I, 12F-G).

Remarks
We propose Fagea bolivari Dresco, 1949 as a junior synonym of Prosalpia martini Simon, 1878. Of the characters mentioned in the original description by Simon, the dorsal black patch on abdominal tergites 2-4 is distinctive, as it never occurs in any other species of Dicranopalpus . Also, the small femoral pedipalpal apophysis of the female holotype (Dresco 1949: 340, fi g. 7) is consistent with that of female Prosalpia martini . Thus, the name Dicranopalpus martini (Simon, 1878) is valid for this taxon. A detailed redescription of D. martini and the related species Dicranopalpus pulchellus is in preparation. (Simon, 1879) Figs 4A, 5A, 6A, F, 7-8, 9B, 10

Diagnosis
Medium-sized species, in both sexes pedipalp patella, tibia and tarsus with dense cover of trichomes. Coxae apically with dark spot.    Dresco, 1953 (ZUPV 4542 4A). Ground colour pale yellowish. Prosoma with groups of small, black spots lateroanteriorly and posteriorly of eye tubercle; dark spots near ozopores. Ozopores small, round. Prosomal tergites V-VI and opisthosomal tergites III-IV pale yellowish brown, darker laterally, following abdominal tergites brown, with paramedian pairs of dark brown triangular patches and light spots, and transverse rows of small dark spots. EYE TUBERCLE. Glossy black, wider than long, positioned at less than its length from anterior margin of carapace. Dorsally with small setae. VENTER AND COXAE. Yellowish, distal coxal margins black, with indistinct central stripe; genital operculum with brown central band; genital operculum, coxae and coxapophyses with long black setae; sternites with small black setae, margins of sternites with dark brown, transverse rows of brown spots.

FRANCE -
CHELICERAE. Pale yellowish, second segment dorsally smoked with brown; segment I with ventral spur; segment I dorsally, and segment II dorsally and medially near cheliceral claw with black setae (Fig. 5A).
PEDIPALPS. Robust compared to other species treated herein (Figs 6A, F, 7A-B); femoral apophysis conspicuous, slightly more than half as long as femur width at its base, pale yellowish, covered with black setae; femur dark, central region lighter, distal part darker. Patella and tibia brown, patella robust, covered with sensory setae as well as trichomes (Fig. 6A); solenidia typically occur grouped or in a row on dorsal side of pedipalp patella (Fig. 7B); patellar apophysis about as long as patella; also tibia with dense cover of trichomes, and with black sensory setae; mediodistal apophysis an indistinct hump (Figs 6A, 7B). Tarsus slightly bent ventrally in distal quarter section; claw with comb-like row of fi ve minute denticles.
DORSUM. Eye tubercle, venter and coxae as in male. Dorsum with paramedian pairs of dark-brown patches.

Distribution and ecology
Dicranopalpus insignipalpis is endemic to Corsica, France (Fig. 10). It is a common mountainous species. According to recent data (JM), its elevational distribution ranges from 300 to 1500 m (13 records), with the majority between 800 m and 1300 m. The upper range may be higher so long running water and adjoining forest cover are present. Generally, D. insignipalpis is a forest-dwelling species, confi ned to forest types offering a minimum of the required air humidity. Consequently, highest population density is found in its upper areas with more abundant precipitation, locally occurring in considerable numbers  (2022) on the banks along mountain streams and rivers, under stones, pebbles and dead wood, often close to the waterline. D. insignipalpis lives at ground level; it was never found on rock faces or tree trunks.
Phenology: according to the known records from June to October, adults and juveniles were collected together. Thus, D. insignipalpis assumingly is eurychronous. The maturity period also cannot be defi ned by altitude. In June and July adults and juveniles were collected syntopically at altitudes of 500-1100 m, while in June and August, only adults were present in localities at altitudes of over 1300 m. Dresco, 1953 (revalidation) Figs 4B, 5B, 6B, G, 9D, 11-13

Dicranopalpus cantabricus
Dicranopalpus cantabricus Dresco, 1953: 147-149 (Fig. 4B). Ground colour pale yellowish. Prosoma with dark brown spots and patches laterally and posteriorly of eye tubercle. Anterior and lateral margins of prosoma pale yellowish. Ozopores small, round. Dark brown saddle narrowing on opisthosomal tergites III-IV, widening on following tergites. Opisthosomal tergites with paramedian pairs of white spots and transverse rows of small dark spots. EYE TUBERCLE. Greyish brown with silvery sheen, shallow, canaliculated (with longitudinal groove), slightly wider than long, and slightly tilted backwards, at about its length from anterior margin of carapace, dorsally with few minute black setae. Eyes surrounded by narrow black ring. VENTER AND COXAE. Uniformly pale yellowish with scattered black setae. CHELICERAE. Uniformly pale yellowish, segment I with distinct ventral spur; segment I dorsally, and segment II dorsally and medially near cheliceral claw with stout black setae (Fig. 5B). PEDIPALPS ( Fig. 11A-C). ( Colouration of specimen faded, colour pattern probably as in D. gallaecicus sp. nov.) Compared to D. gallaecicus sp. nov., segments longer and more slender; femoral apophysis pale, more than half as long as femur width at its base, femur on ventral side with sensilla chaetica that are slightly longer than other setae on femur; patella slender, apophysis about as long as patella, slightly widening in distal half, in lateral view curved upwards (Figs 6G, 11A, C). Mediodistal apophysis of tibia a small knob (Fig. 6B). Tarsus straight, slightly bent ventrally in distal third section; claw pectinate (Fig. 11B).
LEGS. Leg lengths I-IV (in parentheses femur lengths): 25 (4.6); 46 (8.3); 28 (5.0); 37 (6.7). Colouration faded, colour pattern probably as in D. gallaecicus sp. nov. Leg segments cylindrical in cross section, dorsal and dorsolateral sides of femora with sensilla chaetica and a few small black spines. Femur I on dorsodistal half with trichomes, legs II to IV with trichomes, absent only on ventral side proximally. Patellae and tibiae densely covered with trichomes. Tibia II with 5 pseudo-articulations; metatarsi I-IV have 4, 9, 4 and 6 pseudo-articulations, respectively. Numerous bipterate setae on prolateral sides of metatarsus III and IV and in decreasing numbers on proximal tarsomeres 1-10 of leg III and 1-19 of leg IV. PENIS (Fig. 12). Long and slender; length 1.94; truncus widest in basal ⅓, tapering to glans (Fig. 12A). Intrinsic penial muscle in about basal 2 /5. Glans longer than wide, ventral margin continuing in a smooth line to horns (Fig. 12G); glans dorsally and ventrally sclerotized and provided with canaliculi; dorsal cavity large and ovoid (Figs 12C-D, F-G). Dorsal truncus-glans transition in a wide angle. A pair of sensory setae (Fig. 12E). Stylus about as long as glans, slightly curved ventrally; two slender horns positioned on stylus.  PEDIPALPS (Fig. 11D-E). Femoral apophysis robust, nearly as long as femur width at base; patellar apophysis large, reaching to base of tibial apophysis, as thick as tibia. Mediodistal apophysis of tibia stout, as long as wide (Fig. 11E). Femur ventrally with large plumose setae, tibia ventrally with a row of 14 robust plumose setae.

Distribution and ecology
The locality of the redescribed specimens (Camarmeña, Asturias) is about 10 km from the type locality of D. cantabricus (Puertos de Áliva, Cantabria), in the Picos de Europa National Park (Fig. 13). Roewer (1953)

Diagnosis
Medium-sized species, males with dark prosoma, with pale patch in front of eye tubercle. Legs dark brown, femora lack trichomes.

Etymology
The name of the new species refers to the type locality in the north-westernmost Spanish autonomous community of Galicia, in Latin Gallaecia, here with masculine ending, and used as an adjective.    (Figs 4C, 15). Ground colour pale yellowish. Prosoma dark brown to almost black, anterior margin of prosoma, and area proximally of eye tubercle pale yellowish. Ozopores oval-shaped, small. Black saddle wide on prosoma, narrowing on opisthosomal tergites I-IV, widening on following tergites. Opisthosomal tergites with paramedian rows of white spots. Tergite IX and lateral margins of opisthosomal tergites pale. EYE TUBERCLE. Base pale yellowish, dorsally greyish, with silvery sheen most distinct around eyes, shallow, canaliculated, slightly wider than long, and somewhat tilted backwards, at less than its length from anterior margin of carapace, dorsally with a few minute black setae. Eyes with narrow black ring. VENTER AND COXAE. Uniformly pale yellowish, with scattered black setae. CHELICERAE (Fig. 5C). Pale yellowish, segment I with dorsolateral brown patch, segment II in dorsoproximal area dark; ventral spur distinct, segment I dorsally, and segment II dorsally and medially near cheliceral claw with stout black setae.

Material examined
PEDIPALPS (Figs 6C, H, 14A-C). Pale yellowish, femur in proximal half, and patella and tibia dorsally with dark brown patches; femoral apophysis pale, robust, about half as long as femur width at its base, femur on ventral side with sensilla chaetica that are slightly longer than other setae on femur; patella slender, apophysis stout, shorter than patella, rod-shaped, widest in distal half (Fig. 6H), in lateral view distinctly curved upwards (Fig. 14A). Tibia mediodistally with a protrusion (Fig. 6C). Tarsus straight, slightly bent ventrally in distal third section; claw pectinate ( Fig. 14B).
Female LENGTH. 4.3, width of prosoma 2.5, BLI 1.1. DORSUM (Figs 4D, 15). Prosoma with dark brown spots, lateral margins near ozopores dark brown; anterior margin of prosoma and area proximally of eye tubercle silvery. Opisthosoma with small dark brown paramedian patches on tergites III-IV, tergites V-VI with extended dark brown paramedian patches, central area light brown. Opisthosomal tergites with paramedian pairs of white spots and transverse rows of small dark spots. VENTER AND COXAE. Pale yellowish, with scattered black setae. All coxae with subapical dark band. Margins of sternites dark. PEDIPALPS ( Fig. 14D-E). Pale yellowish, with dark brown bands on basal half of femur, distal portion of patellar apophysis and tibial apophysis; tibia dark in midsection, dorsolaterally with dark brown elongated patch. Femoral apophysis robust, half as long as femur width, patellar apophysis large, reaching base of tibial apophysis, as thick as tibia. Mediodistal apophysis on tibia as long as wide. Tarsus straight, claw pectinate.

Distribution and ecology
Dicranopalpus gallaecicus sp. nov. is a Spanish endemic species, its distribution probably being restricted to the Galician region (Fig. 13).
The type locality (Fig. 17A) is a wooded slope consisting of a Pinus L. and Eucalyptus L'Hér. tree plantation, undergrowth mainly with Common gorse ( Ulex europaeus L.). Individuals were present in considerable numbers, and could easily be collected especially on and among shed strips of Eucalyptus bark, hanging from the trees or laying on the forest fl oor. In two cases, individuals were found in resting position on a shaded rock face.
Records based on photos (from https://www.biodiversidadvirtual.org/insectarium/; record number in parentheses) included (Fig. 13 (Figs 4E, 18A). Ground colour pale yellowish brown. Prosoma with dark brown patches and spots, anterior margin pale, lateral margin near ozopores dark brown. Ozopores oval-shaped, small. Saddle dark brown, narrowing on opisthosomal tergites I-IV, widening on following tergites. Opisthosomal tergites with paramedian pairs of white spots. Lateral areas of tergites VI-IX mottled with light and dark spots. EYE TUBERCLE. Dark brown with silvery ring around eyes, canaliculated, slightly wider than long, at about its length from anterior margin of carapace, dorsally with few minute black setae. VENTER AND COXAE. Uniformly pale yellowish brown, with black sensory setae. CHELICERAE (Fig. 5D). Small, smooth, pale yellowish, segment I with ventral spur, segment I dorsally, and segment II dorsally and medially near cheliceral claw with black setae.  Dresco, 1948 (red symbols) and Dicranopalpus catariegensis sp. nov. (blue symbols). Dots, studied samples; squares, literature records. Type localities are indicated by a black spot. The two French sites indicated by '?' were recorded by Dresco (1948) as D. pyrenaeus ; the Spanish '?' was recorded by Bellés (1978) (Fig. 19). Slender, pale yellowish, femur in proximal half, patella and tibia darker; femoral apophysis pale, robust, more than half as long as femur width at its base, femur ventrally and mediodistally with sensilla chaetica; patellar apophysis slender, tapering distally, as long as or longer than patella, in lateral view slightly curved upwards (Fig. 6I). Tibia mediodistally with a small protrusion (Fig. 6D). Tarsus slightly bent ventrally in distal third section; claw pectinate (Fig. 19B).

Female
LENGTH. 3.1, width of prosoma 2.2, BLI 1.0. DORSUM (Fig. 18B). Colours faded in available samples. Prosoma as in male, saddle dark brown, narrowing on opisthosomal tergites I-IV, continuing on following tergites as a broad median band. Opisthosomal tergites with paramedian pairs of white spots, lateral areas mottled with light and dark spots. PEDIPALPS (Fig. 19D-E). Femoral apophysis robust, almost as long as femur width. Long and robust plumose setae on apophysis, and ventral and mediodistal side of femur; patellar apophysis large, with tapering tip, not reaching tibial apophysis, about 4 /5 as thick as tibia. Mediodistal apophysis as long as wide. Tibia ventrally with twelve robust plumose setae.
Phenology: adults probably from late summer to late autumn.

Diagnosis
Medium-sized, with long brown legs. Male: prosoma pale with light brown patches, opisthosoma dark brown.

Etymology
The name of the new species refers to its east-Pyrenean distribution, combining names of the Spanish autonomous community of Cataluña and of the French department Ariège ('cat-arieg-ensis').

Material examined
Holotype  (Fig. 4F). Ground colour pale yellowish. Prosoma pale yellowish, with light brown patches, margins near scent glands and area posteriorly of eye tubercle darkened. Ozopores oval-shaped, small. Dark brown saddle, narrow on opisthosomal tergites III-IV, widening on tergite IV. Opisthosomal tergites with paramedian pairs of white spots and transverse rows of small dark spots. Lateral areas of tergites VI-IX lighter. EYE TUBERCLE. Slightly wider than long, not tilted backwards, canaliculated, at about its length from anterior margin, greyish silvery, midsection dark brown, provided with about ten minute black setae; eyes with narrow black ring. VENTER AND COXAE. Uniformly pale yellowish brown, with black sensory setae. CHELICERAE (Fig. 5E). Pale yellowish, slightly darker dorsally; segment I with ventral spur, segment I dorsally, and segment II dorsally and medially near cheliceral claw with black setae. PEDIPALPS (Fig. 22A, C). Pale yellowish, dorsal sides of femur, patella and distal part of tibia somewhat darker; claw pectinate ( Fig. 22B). Femoral apophysis slightly shorter than wide at its base; patellar apophysis as long as or slightly shorter than patella, curved upwards in lateral view (Fig. 6J); tibia mediodistally with small protrusion (Fig. 6E). Claw pectinate (Fig. 22B).
Female LENGTH. 4.9, width of prosoma 2.5, BLI 1.8. DORSUM (Fig. 18B). Prosoma as in male, saddle consisting of dark brown paramedian patches, narrowing on opisthosomal tergites I-IV with median area light brown. Opisthosomal tergites with paramedian pairs of white spots, lateral areas mottled with light and dark spots and patches, and transverse rows of small dark spots. EYE TUBERCLE. Brown with silvery sheen, lighter around eyes; canaliculated, with about ten minute black setae.

Distribution and ecology
A species endemic to the eastern Pyrenees (Fig. 21). The Montseny, a massiv between Barcelona and Girona provinces where Rambla & Perera (1995) recorded the species as D. martini , seems to be an isolated southern record. Distribution data suggest that D. pyrenaeus and D. catariegensis ranges could meet in the northwestern part of Lleida province. A sample of D. catariegensis sp. nov. from the Hautes-Pyrénées (CHW 467) is within the distribution range of D. pyrenaeus . Most records are from altitudes 1200-1400 m a.s.l.; a high locality is from Espot at 2141 m a.s.l. We found adult specimens mostly on tree trunks, under loose stones and on rock faces. The sex ratio in favour of males (21M, 6F) in our samples suggests that females mostly select daytime shelters at ground level, while males rest higher up, and are thus easier to fi nd. Juveniles live at ground level.
Phenology: insuffi ciently known, adults from June to end of August; juveniles from July to October, suggesting that eggs hatch in summer and autumn, juveniles overwinter and adults appear in spring. 3. Prosoma mottled with brown. Patellar apophysis slender, as long as patella, with tapering tip (Fig. 6J)

Discussion
Our revision of the Dicranopalpus pyrenaeus species group from the Pyrenees and northern Spain, primarily based on penial and pedipalpal morphology, resulted in the redescription of D. pyrenaeus , the revalidation of D. cantabricus and descriptions of two hitherto hidden species, D. gallaecicus sp. nov. and D. catariegensis sp. nov. Also, we propose D. bolivari as a new synonym of D. martini .
The genus Dicranopalpus (superfamily Phalangioidea) presently has no family assignment but is placed in the so-called " Dicranopalpus group", containing seven genera (Crawford 1992;Pinto-da-Rocha & Giribet 2007). It has been provisionally placed in different subfamilies of Phalangiidae by various authors: Phalangiinae (Dresco 1949), Gyantinae (now Gyinae) (Rambla 1975) and Oligolophinae (Rambla 1960), or in Sclerosomatidae (Wijnhoven & Martens 2019). Martens (2021) suggested that the Dicranopalpus group probably has a long evolutionary history in Europe, and may precede the European Journal of Taxonomy 839: 39-73 (2022) Phalangiidae/Sclerosomatidae split. Morphological peculiarities such as the long apophysis on the pedipalpal patella also occur in other genera. For example, in Megistobunus Hansen, 1921 (Phalangiinae) and in some central Asian Gyinae (Sclerosomatidae, e.g., Rongsharia Roewer, 1957) (Martens 1982;Wijnhoven & Prieto 2015). In order to fully resolve this issue, genetic analyses considering both European and central Asian species will be required. In our opinion, the genus Dicranopalpus will probably deserve a separate subfamily or family assignment.
Currently Additionally, the phylogenetic relationships within Dicranopalpus have not been satisfactorily resolved. As mentioned before, on the basis of morphological traits, two distinct species groups can be recognised. These are the sister groups of D. ramosus / caudatus (Wijnhoven & Prieto 2015) and D. martini / pulchellus (Rambla 1975). Relationships among the nine remaining species are unknown. Thus, the D. pyrenaeus species group as defi ned in this study (comprising D. cantabricus , D. gallaecicus sp. nov., D. pyrenaeus and D. catariegensis sp. nov.) may not be monophyletic.
We recommend that further characterisation of species in this genus should focus on detailed descriptions of the pedipalps (shape, lengths and relative dimensions of apophyses; setation) and genital structures (in particular the three-dimensional structure of the penis glans).
In the four Iberian species D. catariegensis sp. nov., D. pyrenaeus , D. cantabricus and D. gallaecicus sp. nov., cheliceral morphology is of minor importance; the dorsal colour pattern is rather similar, while, for unknown reasons, males are darker from east to west.
The mentioned four species occupy largely allopatric distribution areas (Figs 13,21). The distribution of the Pyrenean D. pyrenaeus and D. catariegensis sp. nov. shows distinct similarities with species of the genus Centetostoma Kratochvíl, 1958(Martens 2011, with C. scabriculum (Simon, 1879) being the western and C. juberthiei Martens, 2011 being the eastern Pyrenean element, while C. ventalloi (Mello-Leitão, 1936) overlaps with both species of Dicranopalpus . According to Martens (2011), these three species of Centetostoma may have evolved from isolated populations as a result of Pleistocene climatic changes. D. pyrenaeus and D. catariegensis sp. nov. may share a similar phylogenetic origin.
In the species under study, the dorsal truncus/glans junction is a heavily sclerotized zone, that distally divides into two ridges, surrounding a slit-shaped to oval-shaped cavity (Fig. 3C-D). Inside there is an infl atable and retractable integument. This structure is absent in the D. ramosus / D. caudatus species group, supporting its unique position within the genus. Other species of the genus do possess this glans cavity, with the exception of D. brevipes . Here, this cavity is lacking, possibly due to its ground dwelling habits, imposed by a xeric climate (Wijnhoven et al . 2020: fi g. 5). Interestingly, Mitopus morio (Fabricius, 1799) (Phalangiidae, Oligolophinae) has a similar glans cavity, including a bladder-like structure (e.g., Martens 1978: fi g. 658). Again, genetic analyses will be required to resolve whether this feature has a homologous or analogous origin.
Many questions regarding the biology of Dicranopalpus (including their seasonal phenology, horizontal and vertical distribution, and ecology) remain unanswered. Rambla (1975)