The Centetostoma Scabriculum Complex—A Group of Three Cryptic Species (Arachnida: Opiliones: Nemastomatidae)

The Centetostoma scabriculum complex—a group of three cryptic species (Arachnida: Opiliones: Nemastomatidae)

JOCHEN MARTENS Institute of Zoology, Johannes Gutenberg University Mainz, Department of Systematic Zoology, Müllerweg 6, D-55099 Mainz, Ger- many. E-mail: [email protected]

Abstract

Nemastoma scabriculum Simon, 1879 turned out to be a group of three closely related species of the Pyrenees in SW Eu- rope (France and Spain). Though the species are similar in general habit they can easily be recognized by external morphology (structure of the apophysis of male chelicerae) and male genital morphology (penial structure). For Nemastoma scabriculum Simon, 1879 sensu stricto a lectotype is designated. The synonymous Nemastoma ventalloi Mello-Leitao, 1936 is considered to be a distinct species and is re-described. The third species of the group is described as new, Cente- tostoma juberthiei sp. n. Though partly sympatric in their restricted Pyrenean area, the individual species are found to be small-scale allopatric; only twice have two species been found in syntopy in narrowly circumscribed localities. Centeto- stoma Kratochvíl, 1958 is confined to presently comprise one Southwest Alpine and three Pyrenean species.

Key words: Centetostoma juberthiei, new species, Centetostoma ventalloi, taxonomy, phylogeny, lectotype, synonymy, Pyrenees, France, Spain

Introduction

Taxonomic history. In 1879 Simon described a new nemastomatid species from the central Pyrenees, Nemastoma scabriculum Simon, 1879. In his description he mentioned the patella of the male palp to be armed with a medio- distal spur, much smaller than in Nemastoma dentipalpe Ausserer. Over the decades this species attracted little attention and in general was mentioned only in catalogues. Roewer (1914, 1919, 1923) presented a rough drawing of the dorsal side of the body and mentioned it only from the type locality. Furthermore, in his 1914 publication and in all subsequent publications in which he treated nemastomatids (Roewer 1919, 1923, 1951) he pointed out that he saw Simon’s type series originating from Saint-Sauveur and that, in the 1951 paper, he obtained samples from Simon’s type material. In 1919 Roewer drew attention to Nemastoma scabriculum as –in his opinion– close relative of Nemastoma centetes Simon, 1879. Strangely, this Simon series of N. scabriculum (‘ex type’) is still present in the Roewer collection and, correctly, is labelled Saint Sauveur, the type locality (now in SMF Frankfurt am Main). Mello-Leitao (1936) described a new species undoubtedly closely related to N. scabriculum and named it Nem- astoma ventalloi Mello-Leitao, 1936. He added a detailed drawing of the dorsal side including chelicerae and pedipalp, showing that in the male chelicerae this species has a deeply split apophysis and that the male pedipalp’s patella lacks a spur. These characters could be verified by inspection of two paratypes and newly collected material. Kratochvíl (1958) divided the polytypic and certainly not monophyletic genus Nemastoma into several genera in a Roewerian typological manner based on external morphology. Few of these genera gained wider acceptance, but several are in usage still today. In his analysis N. scabriculum played no role, but he selected Nemastoma centetes as the type species of his new genus Centetostoma Kratochvíl, 1958. In this study, this species of the western Alps is regarded as a close ally of the Pyrenean N. scabriculum group, which we hereby place in the genus Cente- tostoma (see Systematics on genus level). In the following decades the species remained unattractive for taxonomists. Kraus (1961), Rambla & Perera (1989) and Rambla (1998, 2001) published new records from the Pyrenees, the latter authors from Huesca. Kraus

Accepted by A. Schönhofer: 1 Jan. 2011; published: 4 Mar. 2011 35 (1961) tentatively supposed that the taxa scabriculum and ventalloi might be synonymous. This was definitely supported by Rambla (1980) in the first detailed post-Simon account dealing with Nemastoma scabriculum. She checked the original material of Simon in MNHN Paris and the type material of ventalloi and found the specimens to be indistinguishable from scabriculum. She provided a penial drawing of what she called scabriculum but mentioned a certain variability of the penial characters, especially referring to the lateral wings of the truncus penis (“…parfois rétrécies et peu visibles”), but did not present further morphological details, especially of the male chelicerae. In the course of current nemastomatid studies it turned out that N. scabriculum in reality is a composite group, a name under which three species are hidden which all occur in the Pyrenees (Fig. 1). These are very similar according to habit, coloration and external morphology but nevertheless well distinguishable by male chelicerae and male genital morphology characteristics. Species delineating characters and current taxonomy. The three species distinguished here can most easily be characterized in the male sex by cheliceral morphology (see below: ‘cheliceral types’ I, II and III), by penial characters and partly by armature of the male patella. The following morpho-types, i.e. population groups, can be distinguished: ‘Ventalloi’: Type I: Apophysis of male basichelicerite deeply split at medial side (Fig. 6); without spur on palpal patella medio-distal (Figs 19, 23); wings on penial truncus broadest, slender, pointed and slightly directed upwards (Figs 24–29). ‘Scabriculum’: Type II: Apophysis of male basichelicerite moderately excavate at medial side of male basichelicerite (Fig. 8); with a strong spur on palpal patella medio-distal (Fig. 20–21); wings on penial truncus similar to type I, less broad and less pointed, with compact general appearance, wings only rarely directed upwards (Figs 14– 18). ‘Juberthiei’: Type III: Cheliceral apophysis of male basichelicerite smoothly indented at medial side (Fig. 10); with a spur on palpal patella medio-distal (Fig. 22), often indistinctly developed; wings on penial truncus only slightly exposed sideward, rounded, small and much less distinct than in type I and II species (Figs 30–32). When this complexity of the scabriculum group became evident, the question arose which species Simon (1879) actually described and which taxon was named by Mello-Leitao (1936) as ventalloi. To solve this question turned out to be complicated because Simon quite often did not fix type specimens or at least type series and, even worse, united specimens of what he believed to belong to the same species. Due to this attitude even series from different localities were mixed into in a single series (Schönhofer & Martens 2008 for Trogulus Latreille). This often posed major taxonomic problems especially in groups which contain cryptic species, the existence of which was unknown and unexpected at Simon’s time when diagnostic characters in harvestmen were elaborated in less detail. In the Simon collection in MNHN Paris presently no series is labelled “type” or “types” (pers. comm. by M. Judson). Nevertheless, an original series of the Simon collection is deposited in the Roewer collection (SMF RI/ 967, Frankfurt am Main) and contains two species of the scabriculum group, apparently a secondarily composed series (see below). To solve the taxonomic uncertainties, it is indispensable to establish a lectotype for Nemastoma scabriculum and to set forth the identity of Nemastoma ventalloi. According to the rules of the Code (ICZN 1999, § 74.1), lectotypes are to be chosen, so far as available, from the original type series. As only reliable syntype series we treat the series at SMF which Roewer got from Simon as early as in 1914 (probably even earlier). Its origin is given as Saint Sauveur, the published indication of the type locality of N. scabriculum. This syntype series turned out to be a composite one containing two species, C. scabriculum sensu stricto and C. juberthiei sp. n. Systematics on genus level. Nemastomatid generic division is mainly based on genital and cheliceral morphology and presently reflects a relatively high degree of stability. Papers of Kratochvíl (1958, 1959), Šilhavý (1966), Martens (1978, 2006), Staręga (1976) and Gruber (1976, 1979) and Gruber & Martens (1968) contributed much to our understanding. A first, still incomplete molecular-genetic analysis corroborated the present taxonomy and systematics and underlines that these characters are suitable for generic divisions and placement of species (Schönhofer & Martens 2010). Within the nemastomatid fauna of the Iberian Peninsula (including the northern slopes of the Pyrenees) only representatives of five genera occur: Nemastoma C.L. Koch, 1839, Nemastomella Mello-Leitao, 1936, Mitostoma Roewer, 1951, Acromitostoma Roewer, 1951 and Centetostoma Kratochvíl, 1958. Nemastomella species are widespread all over the Iberian Peninsula, are mostly allopatrically distributed and they all have a similar genital morphology (Dresco 1967, Rambla 1968, Prieto 2004). Acromitostoma is restricted to southern Spain and northern Africa and the only presently verified Iberian Mitostoma species is restricted to the northern and southern slopes of the Pyrenees. Nemastoma (in the sense of Gruber & Martens 1968 and Martens 1978) is known only from a

36 · Zootaxa 2783 © 2011 Magnolia Press MARTENS number of localities in the Pyrenees and Cantabric Mountains (Rambla 1980, 1985, 1998, Delfosse & Iorio 2009). Centetostoma and Nemastomella have been separated in current taxonomic use only shortly. Martens (1978) included present Nemastomella species into Centetostoma. At that time the name Nemastomella was not in use because its type species was based on a juvenile specimen. Staręga (1986) revived Nemastomella, and included eleven species together with the type of Centetostoma, Nemastoma centetes, but did not mention scabriculum. Yet, genital morphology of Centetostoma centetes from the Alps and the scabriculum complex from the Pyrenees is similar insofar as all have an alate (winged) truncus penis below the glans and this might be an apomorphy of a monophyletic group. But alate trunci are also present in the two Acromitostoma species of southern Spain (Rambla 1983) and in nemastomatid species of the Balkans, Turkey and the Caucasus (Martens 2006), certainly not related to Iberian groups. Consequently, this character may not be a reliable indicator of relationship in all cases, and mul- tiple parallel evolutionary events are conceivable. On the other hand, the cheliceral apophysis shape of Centeto- stoma species is close to that of Nemastomella and Acromitostoma species, and with respect to dorsal armament the taxon centetes in reality may be a derived Nemastomella. Probably the genera Centetostoma, Acromitostoma and Nemastomella represent a monophyletic Iberian radiation, with C. centetes as a geographic outlier. Their species are clamped by the unique morphology of cheliceral apophyses. To finally solve this uncertainty we have to expect molecular genetic analyses. Here Centetostoma is treated as a genus of its own including four species, and only the Pyrenean species are dealt with in this paper (for C. centetes see Martens 1978: 140–141). Distribution. Even though the scabriculum complex inhabits a relatively small area within the Pyrenean mountain chain, except for the extreme West, the three species accepted here occur largely allopatrically (Fig. 1). Even in the one belt of area overlap, fine-scaled allopatric distribution prevails. Normally, only one species exists at an individual locality. Furthermore, these three species apparently are not homogeneously distributed, being alto- gether absent over larger distances. This may explain the lack of local co-occurrence and mixed colonies. Even large series (up to nearly 50 specimens) always contained only one species except for two instances:

1) Lac des Bouillouses (Pyrenées-Orientales, France) 1♂ of C. ventalloi among 11♂ and 10♀ of C. juberthiei sp. n. 2) Below Col d’Ares (Pyrenées-Orientales, France) 1♂ of C. ventalloi among 26♂ and 18♀ of C. juberthiei sp. n.

FIGURE 1. Pyrenean distribution of Centetostoma scabriculum (squares, blue), C. ventalloi (dots, green) and C. juberthiei sp. n. (triangles, red). Note two sympatric occurrences of C. ventalloi and C. juberthiei sp. n. with overlaying symbols. The western-most overlaying symbols (scabriculum/juberthiei sp. n.) represent the type locality of C. scabriculum (Saint-Sauveur) and denote a false record for C. juberthiei sp. n.

Most of the specimens were collected by hand under stones and wooden debris and by sifting forest soil litter. A few were loaned from public collections and from the collection A. Schönhofer. I was unable to investigate the Centetostoma material of the Muséum National d’Histoire Naturelle in Paris including the Simon collection. Most of the material used is deposited in the working collection of J. Martens in the Institute of Zoology, Mainz Univer- sity, Germany. Original line drawings were produced using a camera lucida attached to a Carl Zeiss study microscope. Mea- surements were taken by means of a micrometer disc attached to a Leitz stereomicroscope. Measurements of penis wing spans were taken from the original drawings. Body length refers to the dorsal scutum magnum only, not to free tergites. Records in the material sections are ordered consecutively from West to East. Abbreviations used: Apo apophysis, Cx coxa, Fe femur, Mt metatarsus, Pt patella, Ta tarsus, Ti tibia, Tr tro- chanter. All measurements are given in mm.

Collection acronyms used:

CAS Collection A. Schönhofer, presently housed at the Institute for Zoology, Mainz University, Germany CJM Collection J. Martens at the Institute for Zoology, Mainz University, Germany MNHN Muséum National d’Histoire Naturelle Paris, France MNRJ Museu Nacional/UFRJ, Rio de Janeiro, RJ, Brazil SMF Naturmuseum and Forschungsinstitut Senckenberg, Frankfurt am M., Germany

Taxonomy

Centetostoma Kratochvíl, 1958

Type species: Nemastoma centetes Simon, 1879 (original designation).

The genus is here defined comprising four species, one from the western Alps (C. centetes) and three from the Pyr- enees (C. scabriculum, C. ventalloi, C. juberthiei sp. n.). Genital morphology (Figs 11–13): Penial truncus straight, at the basis bulb-like enlarged and bifid, thus the two portions of penial muscles confined to one half of the bulb each; truncus slender, more or less parallel-sided, glans symmetrically tapering to the end, opening of the sperm duct at the very tip; below the glans, an alate (wing-like) structure, slender and rounded (in centetes, ventalloi) or strongly extended and pointed sidewards (in scabriculum, juberthiei sp. n.). Apophysis of male chelicerae (Figs 5–10): Well developed, concave medially and (only in the Pyrenean species) split into two parts, a distal and a proximal one (from dorsal view). Pedipalps: Slender, only in two species (scabriculum and juberthiei sp. n.) with a spur on the patella medio- distally. Legs: Short in the Pyrenean species, (relatively) longer and considerably more slender in C. centetes. Armature of the dorsal side (Figs 2–4): Besides a differing granulation covering the scutum, a more or less indistinct tubercle pair is on each area I-V of the scutum; these are strongly developed with additional long thorns only in C. centetes. Coloration. Depending on age brownish to blackish without silvery marks. This holds true also in C. centetes of the Alps. Distribution. One species in the western Alps, three at medium to high altitude in the Pyrenees except for the western-most part (Fig. 1).

Centetostoma scabriculum (Simon, 1879) sensu stricto Figs 1, 3, 7–8, 12, 14–18, 20–21.

Nemastoma scabriculum Simon 1879: 284; Simon 1881: 90; Roewer 1914: 150; Roewer 1919: 169; Roewer 1923: 675; Roewer 1951: 130; Kraus 1961: 344; Rambla 1980: 198.

Material investigated. Lectotype ♂, FRANCE, Dép. Hautes-Pyrenées, Saint-Sauveur (type locality), collector and collecting date unknown, N 42.867° W 0.017° (syntype series SMF RI/967; this series originally comprised also 1♀ C. scabriculum and 1♂ C. juberthiei sp. n. – see below). Paralectotype ♀, FRANCE, Dép. Hautes-Pyrenées, Saint Sauveur (type locality), collector and collecting date unknown, N 42.867° W 0.017° (SMF 61245, from syntype series SMF RI/967). Further material investigated. 1♂, Dép. Pyrénées-Atlantiques, St. Martin, S of Arette, la Pierre, 1900 m, N 42.97° W 0.73°, J. Martens leg. 20.8.1996 (CJM 3807); 2♀, Dép. Pyrénées-Atlantiques, Pic du Midi d'Ossau, 2700–2880 m, N 42.843° W 0.438°, K. Thaler leg. 13.7.1982 (CJM 3917); 1♀, Dép. Pyrenées-Atlantiques, Col du Pourtalet, Ref. Piombie, 2100 m, N 42.8° W 0.416°, K. Thaler leg. 13.7.1982 (CJM 3916); 15♂ 8♀ Dép. Pyrenées- Atlantiques, Col du Pourtalet, 1800 m, N 42.805° W 0.416°, J. Martens leg. 6.9.1967 (CJM 657); 1♂, Dép. Pyrénées-Atlantiques, E of Col d´Aubisque, 1600 m, N 42.996°W 0.341°, W. Schawaller leg. 1.9.1986 (CJM 2684); 1♂ 1♀, Dép. Pyrénées-Atlantiques, Laruns, Arrens, E of Col d'Aubisque, 1600 m, N 42.996° W 0.218°, K. Thaler leg. 14.7.1982 (CJM 3918); 1♂, Dép. Hautes-Pyrénées, Gavarnie, Fagetum, 1400 m, N 42.733° W 0.01°, P. Beron leg. 1.9.1967 (CJM 3699); 22♂ 11♀, Dép. Hautes-Pyrénées, Cirque de Gavarnie, 1600–1650 m, N 42.683° W 0.006°, J. Martens leg. 26.9.1979 (CJM 2594); 2♂6♀, Midi-Pyrénées, Dép. Hautes-Pyrénées, SW Lannemezan, Val d´Estaragne near Lac de Cap-de-Long, 2050–2150 m, N 42.816° E 0.141°, J. Martens leg. 30.8.1978 (CJM 1771); – SPAIN. 7♂ 2♀, Aragón, Prov. Huesca, Parque Nacional de Ordesa-Monte Perdido, close to the town of Torla, 1400 m, Fagus forest, N 42.633° W 0.1°, J. Niethammer leg. 29.5.1972 (CJM 1796); 8♂ 12♀, same locality, 1850 m, Fagus forest, N 42.633° W 0.1°, J. Niethammer leg. 29.5.1972 (CJM 1799); 1♀, same locality, Torla, N 42.633° W 0.1°, J. Niethammer leg. 24.5.1972 (CJM 1801); 1♂ 2♀ (label indication: 2♂3♀), Parque Nacional de Ordesa-Monte Perdido, Refugio de Góriz, [original label: Zentralpyr., Refugio de Golic], N 42.656º W 0.089°, H. Franz leg. 10.8.1955 (SMF 11668). Taxonomy. Within the scabriculum complex there are two species which carry a spur on the palp patella medio-distally, scabriculum sensu stricto and a hitherto undescribed species from the eastern Pyrenees. The scabriculum syntype series contains a male of each of these species that possess this spur. According to the presented material (Fig. 1) both species are strictly allopatric and according to locality a male and the female were selected as representatives of N. scabriculum sensu stricto and the male fixed as a lectotype. The third specimen of the series (1♂) belongs to the species of the eastern Pyrenees and apparently does not originate from the locality indicated, Saint-Sauveur. This species is described below as new based on fresh material (see C. juberthiei sp. n.). Diagnosis. A Pyrenean species of Centetostoma. Apo of male basichelicerite moderately split at medial side (Fig. 8); with a strong spur on palpal patella medio-distally (Figs 20–21); span of wings on penial truncus broad, but less than those of S. ventalloi and with no mensural overlap (Figs 12, 14–18). Measurements. Leg II (♂, ♀ in parentheses): Fe 1.25 (1.25), Pt 0.3 (0.3), Ti 0.9 (0.9), Mt 1.75 (1.7), Ta 1.4 (1.3). Body length: 1.4–1.7, n=10 (1.7–1.9, n=3). Span of truncus wings: 0.14–0.21 (n=5, Figs 14–18). Description. Body (Fig. 3): Body narrowest at front, towards the rear end slightly enlarged laterally, most sig- nificant at rear third of body; entirely black in adults several weeks after moult to adulthood; older adult specimens encrusted with a thin layer of secretion, which – at least after storage in alcohol – is opaque whitish, which more or less camouflages sculpture of the body’s surface. Dorsal scutum (Fig. 3): With a distinct armature of well-defined button-like tubercles, quite large and positioned in only moderately regular rows. Along an imaginary median line from the rear eye tubercle to the rear end of the dorsal scutum there are about 27–30 individual tubercles not touching each other; intermediate space between tubercles about tubercle’s diameter. Rear end of scutum with a saw-like row of blunt longish tubercles. On area I–V of the dorsal scutum one pair each of larger und slightly higher tubercles, the inter-distances enlarging towards the rear end of the scutum.

CENTETOSTOMA SCABRICULUM COMPLEX Zootaxa 2783 © 2011 Magnolia Press · 39 FIGURES 2–4. Dorsal side of body, %. 2: Centetostoma ventalloi (Col de la Core, CJM 752). 3: C. scabriculum (Torla, CJM 1799). 4: C. juberthiei sp. n. (Lac des Bouillouses, CJM 1792).

Tuber oculorum (Fig. 3): Anterior end on the front margin of the dorsal scutum; irregularly armed with two longer (front) to shorter (middle and rear) blunt tubercles. Supracheliceral lamellae (Fig. 3): Split into several longish and upwards-extended comb-like structures, each single tooth armed with a short spiny brush on top. Chelicerae (Figs 7–8): For male only; basal segment dorso-distally with large bulky frontad-directed Apo, markedly broadened at basis, slightly surpassing the dorso-frontal margin of chelicera’s basal segment; no marked incision at the dorsal part of the Apo (medial and lateral view; Fig. 7). In dorsal view, Apo moderately indented at medial side, the curvature forming an approximately right angle, quite obtuse in Fig. 8. In medial view (Fig. 7 right) the top of the Apo rounded, the basal part notably broadened and its medial side deeply carved, protected by a row of strong bristles. Its upper margin is slightly bent to the medial side. Latero-distal side of the basichelicerite and the whole Apo covered with strong bristles (Fig. 7 left). Pedipalp (Figs 20–21): Moderately compact, largest of the three species treated here; Fe slightly enlarged towards distal end, Pt ventrally slightly thickened. Fe to Ta covered with sparse (Fe) to more dense (Pt, Ti, Ta) array of bristles, most of them of the clavate glandular type. There is a distinct medial spur on the distal end of the male Pt which is lacking in the female. Legs: Short in terms of nemastomatid morphology. Fe, Pt and Ti of leg I, III and IV slightly inflated, leg II slender and un-inflated, Fe I from basis to distal end slightly enlarged; Fe, Pt and Ti of all legs with minute evenly distributed acute spines. Pseudoarticulations of femora I–IV (♂, ♀ in parentheses): I 1–2 (1–2), II 5–6 (3–4), III 2–3 (2–4), 3–5 (3–6). Ventral side: Operculum genitale and free sternites with scattered small tubercles, on the sternites only laterally as a short row, on corona analis more densely packed and larger; Cx pro- and retro-laterally with a scattered row of 7–11 blunt tubercles each. Genital morphology (Figs 12, 14–18): Truncus penis extremely thin and slender, at the basis bulb-like inflated and medially incised, the two portions of penial muscles are confined to one half of the bulb each. Truncus distad of the bulb enlargement parallel-sided (Fig. 12). Lower contour of the individual wing is oblique, the upper one mostly horizontal (Figs 14, 18), often with a slight upward turn (Figs 16–17); a downward turn also occurs but probably only artifactually (Fig. 15). Wing span from tip to tip is markedly smaller than in C. ventalloi (Figs 24– 29) with no mensural overlap (see Measurements). Glans is broadest just above the wings (Figs 14, 17) or in the distal half (Figs 15–16, 18), continuously tapering towards the pointed stylus; seminal opening on tip of stylus. Distribution (Fig. 1). Small area in the western part of the French and Spanish side of the Pyrenees. Recently collected material ranges from Pyrenées-Atlantiques (E of Col d’Aubisque: western-most point) to Hautes- Pyrenées (SW Lannemezan, Val d´Estaragne: eastern-most point). The scabriculum type locality, Saint-Sauveur, is

40 · Zootaxa 2783 © 2011 Magnolia Press MARTENS situated in the eastern part of the species’ area. From the Spanish side of the Pyrenees C. scabriculum sensu stricto is only known from the province of Huesca, two localities in the Parque Nacional de Ordesa-Monte Perdido. According to the taxonomy advocated here, these series represent the first verified records for the Spanish fauna. Older unconfirmed records of “scabriculum” refer to Guarda (Portugal; Bacelar 1928), Prieto (2003) does not mentioned specific records of Centetostoma; he just followed the view of Kraus (1961). Rambla & Perera (1989) and Rambla (1998) recorded eleven localities from the province of Huesca, most of them west of Ordesa valley and, consequently, these latter records probably all do refer to C. scabriculum sensu stricto. Presently, the distributional area of C. scabriculum sensu stricto seems to be strictly allopatric to the eastern species C. ventalloi but the western-most locality of the latter species (valley of Neste de Couplan river, SW of Lannemezan) is only a few kil- ometres apart. Ecology. Ten records indicate an altitudinal range from 1400–2800 m (vertical belt 1400 m) and among them five records between 1400–1800 m, three above 2000 m. The species occurs in montane forest, often in open places, under rocks and stones, also in chalky, relatively dry areas (St. Martin, S of Arette). Above the timberline, C. scabriculum lives in open grassland or in nearly bare alpine rock debris under stones. Due to little sampling in high altitude, its upper limit is unknown but may not reach far beyond 2800 m, the highest record for the whole species complex at present.

Centetostoma ventalloi (Mello-Leitao, 1936) Figs 1–2, 5–6, 11, 19, 23–29

Nemastoma ventalloi Mello-Leitao 1936: 9; Kraus 1961: 344; Rambla 1980: 198.

Material investigated. The type and other material originally deposited in the Natural History Museum of Barce- lona are apparently lost (C. Ribera pers. comm. to A. Schönhofer). Paratypes in the Rio de Janeiro Museum of Nat- ural History (see Mello-Leitao 1936: 10) have been inspected. Type material. 1♂ paratype, SPAIN. Prov. Lleida (Lérida), Val d’Aran, D. Ventalló leg., date unknown (MNRJ 42538); 1♂ paratype, same data (MNRJ 42545). Further material investigated. FRANCE. 2♂, Midi-Pyrénées, Dép. Hautes-Pyrénées, SW Lannemezan, above Fabian, valley of Neste de Couplan, 1550 m, N 42.783° E 0.216°, J. Martens leg. 30.8.1978 (CJM 1775); 1♂, Dép. Ariège, near St. Girons, 15km SE of Castillon, Col de la Core, 1400 m, N 42.854° E 1.103°, J. Martens leg. 31.8.1967 (CJM 752); 1♀, same locality, 1400–1450 m, under stones in open pasture, N 42.854° E 1.103°, A. Schönhofer leg. 10.10.2009 (CAS 635); 1♂, Dép. Ariège, S of Couflens, Cirque d´Anglade, 1450 m, N 42.74° E 1.20° J. Martens leg. 19.8.1978 (CJM 1687); 5♂ 1♀, Dép. Ariège, S of Massat, Etang de Lers, in light forest, 1450 m, N 42.883° E 1.333°, J. Martens leg. 22.9.1979 (CJM 2567); 1♂, Dép. Ariège, SW Foix, below Col de Péguère, 1300 m, N 42.916° E 1.383°, J. Martens leg. 17.8.1978 (CJM 1789); 1♂ (together with 11♂ 10♀ C. juberthiei sp. n.; see below), Dép. Pyrénées-Orientales, SW Quillan, Lac des Bouillouses, 1900-2100 m, N 42.716° E 1.983°, J. Martens leg. 20.6.1972 (CJM 6844); 1♂, Dép. Pyrénées-Orientales, Prats-de-Mollo-la-Preste, NE Col d’Ares, border of open pasture and Pinus forest, 1501 m, N 42.375° E 2.456°, A. Schönhofer leg. 2.10.2009 (CAS 597, together with 26♂ 18♀ C. juberthiei sp. n., see below); – SPAIN. 1♂, Cataluña, Prov. Lleida (Lérida), N Pico Maladeta, S Las Bordas (Es Bòrdes), Artiga de Lin, river gorge, 1200 m, N 42.733° E 0.7°, J. & B. Martens leg. 1.9.1984 (CJM 2654); 17♂ 10♀, Cataluña, Prov. Lleida (Lérida), Val d’Aran, Artiga de Lin, 1500-1900 m, N 42.678° E 0.705°, A. Schönhofer leg. 3.-4.10.2009 (CAS 589); 2♂, Cataluña, Prov. Lleida (Lérida), SE Viella, Puerto de Bonaigua, open slope with gravel, grass, 2000 m, N 42.683° E 0.733°, J. & B. Martens leg. 30.8.1984 (CJM 2645); 5♂, same locality, under stones in alpine pasture, 2050-2200 m, N 42.663° E 0.98°, A. Schönhofer leg. 3.10.2009 (CAS 581); 6♂ 2♀, Cataluña, Prov. Lleida (Lérida), SE Viella, SE Puerto de Bonaigua, 1400 m, Abies forest, N 42.663° E 0.981°, J. & B. Martens leg. 29.8.1984 (CJM 2642). Taxonomy. Centetostoma ventalloi was tentatively synonymised with scabriculum by Kraus (1961) which was supported by Rambla (1980). However, based on genital morphology, form of male apophysis of chelicerae and lack of a spur on patella of the palp, C. ventalloi clearly represents a species of its own. Diagnosis. A Pyrenean species of Centetostoma. Apo of male basichelicerite deeply split at medial side (Fig. 6); without spur on palpal patella medio-distally (Figs 19, 23); wings on penial truncus broadest of any species within the genus (Figs 11, 24–29).

CENTETOSTOMA SCABRICULUM COMPLEX Zootaxa 2783 © 2011 Magnolia Press · 41 Measurements (♂, ♀ in parentheses): Leg II: Fe 1.1 (1.1), Pt 0.3 (0.3), Ti 0.8 (0.8), Mt 1.4 (1.4), Ta 1.2 (1.2). Body length: 1.3–1.55, n=3 (1.5–1.7, n=3). Span of truncus wings: 0.22–0.24 (n=6, Figs 24–29). Description. Body (Fig. 2): Body narrows towards front, to the rear end slightly enlarged laterally, most signif- icant at rear third. Entirely black in adults several weeks (to months) after final moult to adulthood; older adult specimens encrusted with a thin layer of secretion which – at least after storage in alcohol – is opaque whitish and more or less camouflages the sculpture of the body’s surface.

FIGURES 5–10. Right ♂ chelicerae. 5, 7, 9: lateral (left column) and medial view (right column); 6, 8, 10: dorsal view. 5–6: Centetostoma ventalloi (Col de la Core, CJM 1792). 7–8: C. scabriculum (Torla, CJM 1799). 9–10: C. juberthiei sp. n. (Lac des Bouillouses, CJM 1792).

42 · Zootaxa 2783 © 2011 Magnolia Press MARTENS FIGURES 11–13. Penes, dorsal view. 11: Centetostoma ventalloi (SW Foix, CJM 1789). 12: C. scabriculum (Col d’Aubisque, CJM 3918). 13: C. juberthiei sp. n. (Col de Port, CJM 1726).

Dorsal scutum (Fig. 2): With a distinct armature of well-defined button-like tubercles, quite large and positioned in only moderately regular rows. Along an imaginary median line from the rear end of the eye mound to the rear end of the dorsal scutum there are about 20 individual tubercles; intermediate space between tubercles less than tubercle’s diameter. Rear end of scutum with a saw-like row of acute longish tubercles, their end slightly rounded; this, too, holds true for the two free tergites. On area I–V of the dorsal scutum one pair each of larger, slightly broader and higher tubercles than the tubercle ground coverage, the inter-distances of these tubercle pairs slightly enlarging towards the rear end of the scutum. Tuber oculorum (Fig. 2): Its anterior end on the front margin of the dorsal scutum; irregularly armed with two longer (antlers-like, front) to shorter (middle and rear) blunt tubercles. Supracheliceral lamellae (Fig. 2): Split into several longish and upwards-extended comb-like structures, each tooth armed with a short spiny brush on top. Chelicerae (Figs 5–6): Male, basal segment dorso-distal with a large frontad-directed knotty Apo (Fig. 5), medio-dorsally (lateral and medial view) somewhat incised forming a shallow saddle (Fig. 5). In dorsal view, this Apo is deeply indented, the curvature forming a narrow triangle separating a distal and a proximal part of the Apo (Fig. 6). In medial view the top of the Apo slightly projects over to the medial side resulting in a deep depression in the inner part of the Apo (Fig. 5 right). The Apo is armed with few spines medially, the lateral and dorso-distal side of the basal segment irregularly armed with short strong spines (Fig. 5). Female: as in the male, but no Apo present on the basal segment.

CENTETOSTOMA SCABRICULUM COMPLEX Zootaxa 2783 © 2011 Magnolia Press · 43 FIGURES 14–18. Centetostoma scabriculum, glandes penis, dorsal view. 14: Torla (CJM 1796). 15: Col d’Aubisque (CJM 3918). 16: Val d’Estaragne (CJM 1771). 17: Col du Pourtalet (CJM 657). 18: Torla (CJM 1799).

Pedipalps (Figs 19, 23): Moderately compact. Fe to Ta covered with sparse (Fe) to more dense (Pt, Ti, Ta) array of bristles, most of them are clavate glandular setae, sparse on Fe and Ti, more dense on Ti and Ta. There is no medial spur on the distal end of Pt in both sexes. Legs: Short in terms of nemastomatid morphology. Fe, Pt and Ti of leg I markedly inflated, less so in legs III and IV, leg II slender and un-inflated, all segments of legs I and III–IV with low and small tubercles causing an irregularly rugged surface. Pseudoarticulations of femora I–IV (♂, ♀ in parentheses): I 0 (0), II 3–4 (3–4), III 2–3 (2), 3– 4 (3–4).

44 · Zootaxa 2783 © 2011 Magnolia Press MARTENS FIGURES 19–23. Pedipalps of ♂, medial view. 19, 23: Centetostoma ventalloi (19: Val d’Aran, MNRJ 42545, paratype; 23: Lac des Bouillouses, CJM 6844). 20–21: C. scabriculum (20: Torla, CJM 1799, 21: Saint Sauveur, SMF 967). 22: C. juberthiei sp. n. (Lac des Bouillouses, CJM 1792).

Ventral side: Operculum genitale and free sternites with scattered small tubercles, in a row on the free tergites, more densely packed on the sternites of the corona analis; coxae pro- and retro-lateral with a scattered row of 6–8 blunt tubercles each. Genital morphology (Figs 11, 24–29): Truncus penis extremely thin and slender, at the basis bulb-like inflated and medially incised. Distad of the truncus bulb narrowed (dorsal/lateral view), from about half length slightly but continuously broadened towards the lateral wings. Lower contour line of the individual wing is oblique, the upper one horizontal with a slight upward turn. In the scabriculum complex wing span is largest in ventalloi (see Measure- ments). Glans is continuously tapering to the pointed stylus, the latter not well differentiated, and to the opening of the seminal duct. Variability: Male genital morphology is subject to some variation. It is most obvious in the truncus wings below the glans: Its span over the lateral tips differs slightly. The single wing may be more compact (Figs 25, 27) or more slender (Figs 26, 29), straight (Figs 24, 29) or curved upwards (Figs 26, 28). There is no overlap in wing span measurements between C. ventalloi and C. scabriculum (see below).

CENTETOSTOMA SCABRICULUM COMPLEX Zootaxa 2783 © 2011 Magnolia Press · 45 FIGURES 24–29. Centetostoma ventalloi, glandes penis, dorsal view. 24: Péguère (CJM 1789). 25: Cirque d’Anglade (CJM 1687). 26: Neste de Couplan (CJM 1775). 27: Val d’Aran (MNRJ 42545, paratype). 28: Val d’Aran (MNRJ 42538, paratype). 29: Col de la Core (CJM 752).

46 · Zootaxa 2783 © 2011 Magnolia Press MARTENS Distribution (Fig. 1). Type locality is in the Spanish Province of Lérida, in the Vallée d’Aran (Val d’Aran), from where the species was recently re-collected (see Material). Further fresh material originates from a relatively small area on the French side of the Pyrenees from the départements of Hautes-Pyrénées (western-most record: SW Lannemezan, valley of Neste de Couplan river) to Ariège and Pyrénées-Orientales (eastern-most record: Lac des Bouillouses, SW Quillan). Most localities originate from Ariège, where the species apparently has its strongholds. There is a considerable area overlap with C. juberthiei sp. n. Ecology. Eight records for altitudinal distribution range from 1200–2000 m (vertical belt 800 m), among them five from 1400–1550 m. Most records originate from light forests and clearings in forests, under stones and decay- ing wood. The only locality from 2000 m is from a rock-covered slope with low grass and herbs. Adults appear all year round and regularly overwinter. At one occasion (CAS 589) a number of specimens were found walking freely on mossy upper parts of large beech tree roots at night (A. Schönhofer pers. comm.).

Centetostoma juberthiei sp. n. Figs 1, 4, 9–10, 13, 22, 30–32.

Holotype ♂ FRANCE. Dép. Pyrénées-Orientales, SW Quillan, Lac des Bouillouses, 1900–2100 m, N 42.716° E 1.983°, J. Martens leg. 20.6.1972 (CJM 7004). Paratypes 10♂ 10♀; same data as holotype (CJM 1792; originally together with 1♂ C. ventalloi, see above); 26♂ 18♀, Dép. Pyrénées-Orientales, Prats-de-Mollo-la-Preste, NE Col d’Ares, border of open pasture and Pinus forest, 1501 m, N 42.375° E 2.456°, A. Schönhofer leg. 2.10.2009 (CAS 576, together with 1♂ C. ventalloi, see above). Further material investigated. FRANCE. 1♂ from syntype series C. scabriculum SMF RI/967, now in SMF 61246; this series originally comprised also 1♂ 1♀ C. scabriculum, see above); 1♂, Dép. Ariège, 30 km S St.Girons, Port d'Aula (road D703), 1500-1500 m, N 2.456° E 1.116°, J. Martens leg. 3.9.1967 (CJM 782); 2♂ 2♀, Midi-Pyrénées, Dép. Ariège, SE of Col de Port, ESE of St. Girons, Forêt de Sanzet, 1200–1300 m, N 42.896° E 1.449°, J. Martens leg. 26.8.1978 (CJM 1726); 2♀, Dép. Pyrénées-Orientales, above Mt. Louis, Tet valley, Kiefern- wald, 1700 m, N 42.506° E 2.1°, W. Schawaller leg. 28.8.1980 (CJM 2672); 1♂ 1♀, Dép. Pyrénées-Orientales, 28 km W of Céret, Pic de Costabonne, 2400–2465 m, alpine grassland, N 42.416° E 2.466°, J. Martens leg. 25.8.1997 (CJM 3633); 6♂ 1♀, Dép. Pyrénées-Orientales, below Col d’Ares, grass, small brook, below stones, 1500 m, N 42.366° E 2.45°, J. & B. Martens leg. 26.8.1984 (CJM 2626); 1♀, Dép. Pyrénées-Orientales, Mont Canigou, high alpine habitat, appr. N 42.519° E 2.457°, H. Franz leg. 25.8.1953 (SMF 11728/1); 2♀, Dép. Pyrénées-Orientales, Mont Canigou, 2200–2500 m, appr. N 42.519° E 2.457°, H. Franz leg. 25.8.1953 (SMF 11749/2); 1♂, Dép. Pyrenées-Orientales, Massif du Canigou, Gomp de Cady, 2400 m, N 42.516° E 2.459°, K. Thaler leg. 25.8.1982 (CJM 3919); 1♀, Dép. Pyrénées-Orientales, Haut Vallespir, NW Prats-de-Mollo, 1600 m, N 42.409° E 2.475°, J. Martens leg. 16.7.1994 (CJM 3895); 1♂ 1♀, Dép. Pyrénées-Orientales, Tech valley, SW of Amélie-les-Bains, la Preste, 1300 m, N 42,4° E 2.488°, J. Martens leg. 23.8.1997 (CJM 3624); 1♂ 2♀, Dép. Pyrénées-Orientales, between Ax-les-Thermes and Querigut, 1150 m, N 42.708° E 2.55°, J. Martens leg. 12.8.1993 (CJM 3422); 1♂, Dép. Pyrénées-Orientales, Haut Vallespir, above Serralongue, 950–1100 m, N 42.383° E 2.55°, J. Martens leg. 1.– 30.8.1993 (CJM 3410). Taxonomy. Nemastoma scabriculum Simon, 1879 turned out to be a composite entity embracing three distinct species taxa. Two of this group are already named (scabriculum Simon, 1879, ventalloi Mello-Leitao, 1936; see above), and the third one is described here as a new species. Diagnosis. A Pyrenean species of Centetostoma. Apo of male basichelicerite smoothly indented at medial side (Fig. 9 right); with a small spur on palpal patella medio-distal (Fig. 19), wings on penial truncus only slightly exposed sideward, rounded, small and much less distinct than in C. scabriculum and C. ventalloi (Figs 30–32). Measurements. Leg II (♂, ♀ in parentheses): Fe 1.05 (1.1), Pt 0.3 (0.4), Ti 0.8 (0.8), Mt 1.25 (1.3), Ta 1.15 (1.25). Body length: 1.4–1.7, n=11 (1.6–1.75, n=8). Span of truncus wings: 0.08–0.09 (n=3, Figs 30–32). Description. Body (Fig. 4): Body narrowest at front, towards the rear end slightly enlarged laterally, most sig- nificant at rear third. Entirely black in adults several weeks after moult to adulthood; older adult specimens are encrusted with a thin layer of secretion which – at least after storage in alcohol – is opaque whitish and more or less camouflages sculpture of the body’s surface.

CENTETOSTOMA SCABRICULUM COMPLEX Zootaxa 2783 © 2011 Magnolia Press · 47 Dorsal scutum (Fig. 4): With a distinct armature of well-defined button (lens)-like tubercles, quite large and positioned in only moderately regular rows. Along an imaginary median line from the rear end of the eye tubercle to the rear end of the dorsal scutum there are about 23–25 individual tubercles not touching each other; intermediate space between tubercles slightly smaller than tubercle’s diameter. Rear end of scutum with a saw-like row of acute tubercles, considerably longer than their diameter. On areae I-V of the dorsal scutum one pair each of larger und slightly higher tubercles than the general surface granulation, the inter-distances enlarging towards the rear end of the scutum. Tuber oculorum (Fig. 4): Its anterior end at the front margin of the dorsal scutum; irregularly armed with two longer (front) to shorter (middle and rear) blunt tubercles, slightly broadened distally. Supracheliceral lamellae (Fig. 4): Split into several longish and upwards-extended comb-like structures, each tooth armed with a short spiny brush on top. Chelicerae (Figs 9–10): Male; basal segment dorso-distally with a large rounded frontally directed Apo, extending considerably from the upper side of the basal segment, beyond the front margin of the basis (Fig. 9 right), in medial/lateral view egg-shaped and continuously rounded dorsally (Fig. 9 left); in dorsal view (Fig. 10) the Apo is slightly indented from medial side forming a shallow depression, sometimes even less pronounced than in Fig. 10. Top of the Apo is only rounded, thus a medially overhanging part of Apo is lacking (Fig. 9 right). Slight armature with long bristles on top and medial side of Apo, moderately strong bristles on the lateral side of the basal segment’s distal half. Pedipalp (Fig. 22): Moderately compact, covered with sparse (Fe) to more dense (Pt, Ti, Ta) coverage of bristles, most of them of the clavate glandular type. There is an indistinct medial spur on the distal end of the male Pt. Being opaque it is often difficult to see and can even be absent in rare cases (CJM 3422). The spur is not present in the female. Legs: Short in terms of nemastomatid morphology. Fe, Pt and Ti of legs I, III and IV slightly inflated, in legs III and IV mostly at distal end, leg II slender and un-inflated, Fe II from basis to distal end slightly enlarged; Fe, Pt and Ti of all legs armed with warty irregular tubercles, producing a rugged surface. Pseudoarticulations of femora I–IV (♂, ♀ in parentheses): I 0–1 (1), II 4–5 (3–5), III 2–3 (2–3), 3–5 (3–5). Ventral side: Operculum genitale shiny, tuberculation scanty, row of tubercles on free sternites only laterally; larger and higher, thus more accentuated, tubercles on corona analis. Cx I-IV pro- and retro-lateral with a scattered row of 8-10 blunt tubercles each. Genital morphology (Figs 13, 30–32): Truncus penis extremely thin and slender (most pronounced of the three species treated here), at the basis bulb-like inflated (this part longest of the three species treated here) and medially incised, each lobe with a bundle of penial muscles. Truncus distad of the bulb enlargement most slender (dorsal/lateral view), from there slightly but continuously enlarged to the lateral wings into which the lateral contours turn (Fig. 13). Lower contour is thus quite steep, upper slightly rounded and the wing-like structure is only small. These wings form a shallow pocket. Glans (Figs 30–32) continuously tapering, then slightly enlarging to distal half, again tapering to the pointed stylus, which is not well differentiated. Opening of seminal duct on top of stylus. Variability: Genital morphology quite stable, especially the alate structures below the glans (Figs 30–32). Distribution (Fig. 1). All localities are situated in the central and eastern-most part of the Pyrenees in the départments of Ariège (two localities, the western-most one: Port d'Aula, 30 km S St.Girons) and Pyrénées-Orien- tales (the eastern-most above Serralongue and between Ax-les-Thermes and Querigut). There is a noticeable distributional overlap between C. ventalloi and C. juberthiei sp. n. in the eastern Pyrenees, just northwest and northeast of Andorra, but only in two localities have both species been found in small-scale syntopy, only a single male of C. ventalloi in large series of C. juberthiei sp. n. (Col d’Ares; Lac des Bouillouses) in both. Apparently, there are no mixed colonies comprising both species in approximately equal numbers. For an alleged record of C. juberthiei sp. n. in Saint-Sauveur (Hautes-Pyrenées), see above (C. scabriculum). Ecology. Eleven records for altitudinal distribution range from 1000–2450 m (vertical belt 1450 m), among them four records between 1500–1700 m, two above 2000 m. Forested areas are commonly inhabited, mainly sparsely overgrown places with low vegetation; often under stones. Due to little sampling at high altitude, the upper limit is not well documented. Derivatio nominis. This well discernible but for long overlooked species is dedicated to M. le Professeur Dr Christian Juberthie, former Director of the Laboratoire souterrain du CNRS at Moulis, Ariège, France. Dr. Juberthie is famous for his fundamental and long-lasting studies on biology, ecology and anatomy of various groups of harvestmen including European cave fauna.

48 · Zootaxa 2783 © 2011 Magnolia Press MARTENS FIGURES 30–32. Centetostoma juberthiei sp. n., glandes penis, dorsal view. 30: Lac des Bouillouses (CJM 1792). 31: Saint Sauveur (SMF 967, lectotype C. scabriculum). 32: SE Col de Port (CJM 1726).

Discussion

The Pyrenean Centetostoma complex comprises three closely related and very similar species which occupy largely allopatric distributional areas, though between two species slight area overlap occurs. However, distribution of individual species is irregular and apparently not evenly dispersed. Even in the overlap area, syntopic occurrences have been found only twice, with one of the concurrent species prevailing markedly. From the viewpoint of phylogeny, these three species apparently are quite young and may have been influenced by Pleistocene climatic changes which caused local isolation of populations. Small post-glacial area expansions certainly occurred and led to the present distributions. Climatic conditions along the Pyrenean chain also may have influenced present distribution. There is a marked climatic gradient from the precipitation-rich Atlantic western Pyrenean parts towards the markedly drier Mediterranean eastern parts. These differences may have prohibited a more eastern and western area expansion of the species, respectively. The final generic affiliation of west alpine (C. centetes) and the three Pyrenean Centetostoma species is still unsettled. Similarities in male genitalia between C. centetes and the Pyrenean species, especially C. juberthiei sp. n., point to close relationships of C. centetes and the three latter species. However, shape of male cheliceral apophyses and dorsal armament of the scutum unites C. centetes with the vast number of Nemastomella Mello-Leitao, 1936 species which are widespread in the Iberian Peninsula including the Pyrenees. Alate penis constructions bear- ing lateral “wings” below the glans, unique to C. centetes and the Pyrenean species within the Centetostoma/Acro- mitostoma complex, are structures to fix the two tendons which originate in the two muscle bundles of the penis truncus base. Consequently, such structures may have evolved convergently several times within nemastomatids and a priori cannot be considered hints for close relationships (details in Martens 2006). We may expect more details of relationships to be revealed by molecular phylogenetic analysis.

My hearty thanks are due to A. Kury for loan of paratypes of Nemastoma ventalloi housed in the MNRJ Rio de Janeiro, to P. Beron, W. Schawaller, A. Schönhofer and the late K. Thaler for additional material, to A. Schönhofer for profound technical help, to K. Rehbinder for the drawings of the dorsal side of specimens and chelicerae, to M. Judson for information on the Simon collection in MNHN Paris and to P. Jäger for loan of material housed in the opilionid collection of SMF Frankfurt am Main. Comments from three reviewers (J. Gruber, C. Prieto, A. Schön- hofer) much improved the clarity of this contribution. Feldbausch Foundation and Wagner Foundation at Fachbe- reich Biologie, Mainz University, granted travel funds for field research.

References

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