European Journal of Taxonomy 216: 1-35

http://dx.doi.org/10.5852/ejt.2016.216

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ISSN 2118-9773

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2016 • Martens J. & Schonhofer A.L.

Research article

urn:lsid:zoobank.org:pub:2F526459-E23A-458B-9829-31F4ClBC6C46

The Leiobunum rupestre species group: resolving the taxonomy of

four widespread European taxa (Opiliones: Sclerosomatidae)

Jochen MARTENS 1 -* & Axel L. SCHONHOFER 2

12 Johannes Gutenberg-Universitat Mainz, Institut fiir Zoologie, 55099 Mainz, Germany

* Corresponding author: martens@uni-mainz de

2 Email: [email protected]

1 um:lsid:zoobank.org:author:F01C78Fl-5EBF-48E8-B37E-634DF00E0237

2 um:lsid:zoobank.org:author:91B2971C-D8CE-4F98-903F-C963E83E8C7C

Abstract. Within the central European opilionid fauna the widely used species names Leiobunum rupestre

Herbst, 1799 and Leiobunum tisciae Avram, 1968 pose taxonomic and distributional problems. In addition,

Nelima apenninica Martens, 1969 is close to L. tisciae in terms of external and genital morphology, but

is specifically distinct. While coxal denticulation is largely lacking in N. apenninica , the validity of

the genus Nelima Roewer, 1910 is questioned again. In addition, Leiobunum subalpinum Komposch,

1998, a recently described novelty from the eastern Alps, is closely related to L. rupestre. The four

species are combined as the morphologically defined Leiobunum rupestre species group. Except for

L. subalpinum , they were found to be allopatrically distributed from the Carpathians across central and

Northwest Europe to the south-western Alps. The latter species is locally sympatric and partly

elevationally parapatric to L. rupestre. Leiobunum tisciae is a recently introduced name and here

recognized as a junior synonym of a number of taxa described much earlier, of which L. gracile Thorell,

1876 is re-introduced as oldest available name. Detailed morphological and distributional data for all

taxa are presented.

Keywords. Taxonomy, synonymy, Leiobunum, Nelima, Europe.

Martens J. & Schonhofer A.L. 2016. The Leiobunum rupestre species group: resolving the taxonomy of four

widespread European taxa (Opiliones: Sclerosomatidae). European Journal of Taxonomy 216: 1-35. http://dx.doi.

org/10.5852/eit.2016.216

Introduction

A common problem with the Sclerosomatidae Simon, 1879 (harvestmen) is their morphological plasticity

as well as their general uniformity, making it often difficult to draw well-recognizable species borders.

Even more problematic, especially in the tropics, is the differentiation of higher taxonomic levels from

species category upwards (Hedin et al. 2012). Applying this difficulty to widespread, closely related and

morphologically similar taxa can further hinder discrimination. Here the similar species Leiobunum

rupestre (Herbst, 1799), L. subalpinum Komposch, 1998, L. tisciae Avram, 1968 and Nelima apenninica

Martens, 1969 are investigated and summarized as the Leiobunum rupestre species group, though being

European Journal of Taxonomy 216 : 1-35 ( 2016 )

generally grouped with two different genera. The four species occupy large parts of central and south¬

eastern Europe, parts of Scandinavia (Finland, Norway, Sweden), locally Great Britain, continental Italy

and south-western France, but sympatric occurrences are unknown (Fig. 2), except for L. subalpinum

whose distribution is nested within the area of L. rupestre. Discrimination of L. rupestre and L. tisciae has

been under debate since Martens (1978) included L. tisciae into the central European fauna, even though

male genital morphology was properly described and is straightforward. A number of European countries

stated the presence of one or two of these species thereafter (e.g., Great Britain: Davidson 2009; Den-

_ v

mark: Enghoff 1988; Czech Republic and Slovak Republic: Silhavy 1981; Italy: Chemini 1980; Marcel-

lino 1971; Poland: Star^ga 2004; Norway: Stol 2003, 2010a, b; Estonia: Star^ga 1978; Tomasson et al.

2014; Latvia: Tumss 1963; Spungis 2008), but a complex morphological and taxonomic situation was

evident.

The description of L. tisciae by Avram (1968) was shortly followed by that of Nelima apenninica by

Martens (1969), who soon recognized that both species are fairly similar in terms of genital morphology,

but he was primarily concerned to discriminate Leiobunum rupestre against L. tisciae (Martens 1978).

Up to today, N. apenninica remains poorly known, being recorded from widely scattered localities.

These four species raise taxonomic problems and need to be well defined. L. tisciae and N. apenninica

having been described relatively recently, and especially because L. tisciae spans a large European area,

making it likely that it has been described under an older name before. L. subalpinum , a novelty from

the eastern Alps, added a further complexity to species recognition in this group of locally common

long-legged harvestmen. Taking this argumentation into account and based on representative reference

material, we here revalidate the species status, synonymy and distribution of the representatives of the

L. rupestre species group. The taxa Leiobunum nigripalpe Simon, 1879 and “ Leiobunum sp.”, the latter a

recent immigrant to western central Europe (Wijnhoven et al. 2006), are not part of the L. rupestre

species group and are not treated here.

Material and methods

Material investigated: 54 series of L. rupestre , 40 series of L. tisciae and 20 series of A. apenninica were

checked; details are listed in the Material sections (see Appendix). Also literature sources, which we feel

present reliable records, are listed there.

Collection acronyms as used:

SMNH

AXLS

CIK

CJM

SMF

Naturhistoriska riksmuseet, Stockholm, Sweden

Coll. Axel Schonhofer, Ma in z, Germany

Coll. Ivo Karaman, Novi Sad, Serbia

Coll. Jochen Martens, Ma in z, Germany

Arachnology Section, Senckenberg Museum, Gesellschaft fur Naturforschung, Frankfurt

am Main, Germany

Abbreviations used in the descriptive parts:

Abd, abd

Ceph

Palp

Terg

thorac

Abdomen, abdominal

cephalothorax

femur

metatarsus

palpus, pedipalpus

patella

tarsus

tergit

thoracic

MARTENS J. & SCHONHOFER A.L., The Leiobunum rupestre group

Ti = tibia

To oc = tuber oculorum, ocularium, eye mound

Original line drawings were produced using a camera lucida attached to a Eeitz dissecting Microscope

and a Eeitz Laborlux Microscope. Measurements were taken by means of a micrometer disc using the

same optical devices. All measurements are given in mm.

Distributional data

From a total of 2600 text-recognized pdf files of opilionid literature, those matching the search strings

‘apenninica’, ‘rupestre’ or ‘tisciae’were selected. Following our established synonymy further, references

were extracted manually and all identifiable distributional records of the L. rupestre group were geo-

referenced. Of these records only a limited number of the recorded specimens was available to us and

geographic reconstructions relied on our representative samplings and sound publication data. We

assume Silhavy (1981) and Stargga (1976, 2004) to have correctly discriminated L. rupestre and

L. ‘ tisciae ’ (L. glabrum in Silhavy 1981) and their data already demarked a North-to-South running

borderline between the two species in the Czech Republic and Slovakia (Silhavy 1981) and in Poland

(Stargga 2004). We further interpreted all records east of this line (e.g., L. rupestre in Stargga 1978) and

north of the central mountain ranges in Germany (e.g., Stol 2010a, b) as L. ‘ tisciae ’, primarily based on

investigation of available material. If we found coverage of records sufficiently reliable and indicative

of general distributional interpretation we did not map additional records in the relevant area (for these,

see in more detail: L. tisciae'. Denmark: Meinertz 1964; Enghoff 1988; Great Britain: British

Arachnological Society 2016; L. rupestre and tisciae combined: Germany: Staudt 2016). All reconstructed

localities are given in Fig. 2 and Appendix 1. Up to-date distributional data of Leiobunum subalpinum

was available from Komposch (1998, 2009b) and we did not search for any additional records.

Results

Class Arachnida Cuvier, 1812

Order Opiliones Sundevall, 1833

Suborder Palpatores Thorell, 1876

Family Sclerosomatidae Simon, 1879

Subfamily Leiobuninae Banks, 1893

Genus Leiobunum C.L. Koch, 1839

Type species

Designation by Thorell, 1876: OpiliofasciatusYLerbst , 1798 = Leiobunum rotundum (Latreille, 1798).

Remark

Of the representative material investigated, we found the four hypothesized species well separable upon

a number of somatic and male genital morphological characters, which are summarized in Table 1 for

convenient use. Species appear largely allopatric, except for L. subalpinum , which is firmly nested within

the area of L. rupestre. In addition, parapatric and locally sympatric occurrences of two species under

discussion in East-Central Europe cannot be excluded, especially in South Poland, the central Alps and

close to the Czech/Slovak border area (Fig. 2).

Leiobunum subalpinum Komposch, 1998 is considered most closely related to L. rupestre (Komposch

1998) and, as such, is also regarded as a member of the L. rupestre species group. As it is easily

discriminated from all discussed forms by its dark coxal markings and, apart from L. rupestre , by its

restricted alpine range, it is not featured here (see Komposch 1998; Komposch & Gruber 2004 for

European Journal of Taxonomy 216 : 1-35 ( 2016 )

Table 1. Comparison of somatic characters of Leiobimum apenninicum (Martens, 1969), L. gracile

Thorell, 1876 and L. rupestre Herbst, 1799. L. subalpinum Komposch, 1998 is not included, because

it is very similar to L. rupestre and is easily identified by its black coxal markings. For comparison

with L. rupestre we refer to Komposch (1998). If not otherwise mentioned, characters apply to males

only. Characters outlining a single species are given in bold, more variable characters are given in

italics. The combination of several characters for determination is advised. Palpal spination is variable,

apparently the appearance in ordered rows is an individual character. Body coloration varies with age

and population, aged specimens are darkest.

L. rupestre Herbst, 1799 L. gracile Thorell, 1876 L. apenninicum (Martens, 1969)

in both sexes dorsum smooth,

granulation cannot be

distinguished with 250x, with

a few pointed granules present

palp-femur with irregular rows

of medium-sized triangular

spines on ventral side; dorsal

side few on distal part, ventral

spines more slender, many

pointing downward

palp-tibia slightly concave in

lateral view at ventro-distal end,

therefore forming a flat S-bend

palp-patella appears longer and

less massive

palp-tarsus bent over its whole

length

denticles and hairs on ventral

side of palp-tibia darker and

therefore well visible

eye mound in both sexes always

without spines

in both sexes dorsal coloration

without pair of white spots on

each area along the median line

in females no lateral coloration

of opisthosoma with a blurred

parallel pattern of zigzag lines

in both sexes dorsal granulation

consisting of flat, pointed

granules, regularly spaced, well

visible at 25Ox

palp-femur with irregular rows

of large triangular spines on

ventral side; dorsal side few on

distal part, ventral spines

broader, not so slender

palp-tibia slightly concave at

ventro-distal end, therefore

forming a flat S-bend

palp-patella appears longer and

less massive

palp-tarsus mostly straight,

bent only in distal third

spines and hairs on ventral side

of palp-tibia lighter and

therefore hard to be seen

eye mound in both sexes

without spines, sometimes one

spine present

in both sexes dorsal coloration

with small white spots on each

area along the median line

in females lateral coloration

of opisthosoma with a blurred

parallel pattern of zigzag lines,

best seen at low magnification

in both sexes dorsal granulation

consisting of flat, pointed

granules, regularly spaced, well

visible at 250x

palp-femur with few spines,

normally restricted to ventro-

distal, rarely few large spines

on ventral side, only single

spines on dorso-distal side

palp-tibia not concave at

ventro-distal end, therefore

forming a flat C-bend

palp-patella appears shorter

and more massive

palp-tarsus bent over its whole

length

spines and hairs on ventral side

of palp-tibia lighter and therefore

hard to be seen

eye mound in both sexes with

1-3 small spines on each side,

rarely absent

in both sexes dorsal coloration

with pair of white spots on each

area along the median line

in females lateral coloration

of opisthosoma with a blurred

parallel pattern of zigzag lines,

best seen at low magnification

MARTENS J. & SCHONHOFER A.L., The Leiobunum rupestre group

details) except for general remarks; its geographic range is indicated in Fig. 2. In addition, the poorly

known Leiobunum nigripalpe Simon, 1889, described from the western Alps (France), was

reinvestigated. However, the type series represents a composite collection of at least two different small

Nelima Roewer species, both unaffiliated to the L. rupestre species group. This holds true, too, for the

specimen depicted as Leiobunum nigripalpe in Martens (1978), representing a yet to be identified

Leiobunum species.

Species accounts of the Leiobunum rupestre group

Leiobunum rupestre (Herbst, 1799)

Figs 1A-B, 2, 3A-B, 4A-B, H-G, 5A-C

Phalangium bicolor Fabricius, 1793: 429. The name was synonymized with Gy as annulatus by Latreille

(1804), which was missed by several subsequent authors; some of these used it as valid name, in syn¬

onymy of, or in relation to, L. rupestre (e.g., C.E. Koch 1847, 16: 56; redescription).

Opilio rupestris Herbst, 1799: 4, fig. 1 (material from the type locality Sachsen, i.e., Saxony, examined).

Leiobunum ovale C.L. Koch, 1848: 59, fig. 1540.

Liobunum glabrum L. Koch, 1869: 4, 6-7.

Leiobunum ovale - Roewer 1910: 203. — Silhavy 1956: 174. —Novak & Gruber 2000: 299.

Phalangium bicolor - Simon 1879: 181. —Roewer 1910: 203 (both also listing Phalangium annulatum

in synonymy). —Eessert 1917: 16.

Liobunum rupestre- Simon 1879: 181. — Cantoni 1882: 194. —Hansen 1884: 500. —Kraepelin 1896:

222. —Roewer 1910: 197, 203. —Lessert 1917: 15. —Roewer 1923: 890 (partim). —Hadzi 1931:

145. — Silhavy 1956: 174, figs 437M43.

Liobunus rupestris - Muller & Schenkel 1895: 819.

Liobunum glabrum - Simon 1879: 181.

Nelima glabra - Roewer 1910: 239. —Hadzi 1931: 148.

Leiobunum rupestre - Silhavy 1948: 9, 24, table 6, figs 4-6. — Martens 1978: 408M12, figs 778-780,

788. — Komposch 1998: 26-27, 34, figs 5, 10, 14, 21.

Taxonomic history

At first glance rather complex, but many distributional records from East central, Northwest and North

Europe in reality refer to Leiobunum ‘ tisciae ’ (see below, L. gracile). L. rupestre is a montane species

and well known from low to mid-altitudes of the central and eastern Alps and mountainous areas in the

Czech Republic and north of the Alps in Germany. Its genital morphology is quite characteristic and was

first reliably depicted by Silhavy (1948, figs 4-6; 1956, figs 437-439), fully in accordance with drawings

in Martens (1978, fig. 778) and Komposch (1998, figs 9-10). Leiobunum populations in the Carpathians,

Poland (Stargga 1976), southern Finland (Heinajoki 1944), along the Baltic coast hinterland and Den¬

mark were also largely affiliated to L. rupestre (Martens 1978, fig. 787), though erroneously. These older

determinations mostly neglected the genital morphology of these crucial northern populations, which

was reconsidered after Avram (1968) published a new species, L. tisciae , clearly distinct from rupestre

but similar in external morphology. Due to its wide distribution, the majority of available names for the

eastern and northern populations can be assigned to the synonym of its oldest name L. gracile (see be¬

low).

Excluding the complex synonymy of L. tisciae {gracile), only one uncertain affiliation remains for

L. rupestre : the name Liobunum glabrum L. Koch, 1869. Silhavy (1956: fig. 443) originally treated

‘glabra ’ as the juvenile L. rupestre , but later used this name to distinguish allopatric populations of

western L. rupestre and eastern L. ‘ glabra ’ (later named L. tisciae) in former Czechoslovakia (Silhavy

European Journal of Taxonomy 216: 1-35 (2016)

1981). This was contradicted by Star^ga (2004), because the type locality of L. glabrum , Meran

(northern Italy, southern Alps), is situated deeply within the territory of L. rupestre (Martens 1978).

We suggest following Hadzi (1931) and Martens (1978) to correctly place Leiobunum glabrum in the

synonymy of L. rupestre.

Diagnosis

A medium-sized Leiobunum species with blackish upper side, except for broad, white markings on Ceph

disto-laterally in male (Figs 1A, 3 A), broad, blackish irregular saddle-like marking all over the length of

the body in female with extended white markings laterally from Ceph to abd Area II and a cross stripe

all over area V (Figs IB, 3B). In both sexes series of para-median lines of small white spots on the abd

areae I-V are absent. Contrasting yellowish underside including coxae of all appendages. Wings of

truncus penis broad, broadest in lower third part of truncus (Fig. 5A-B). Coxa IV with retrolateral row

of granules.

Fig. 1. Habit of Leiobunum rupestre species group. A-B. Leiobunum rupestre Herbst, 1799, Slovenia,

Pohorje Mountains, resting at rock faces. A. $. B. §. C-D. Leiobunum apenninicum (Martens, 1969),

Italy, Monesi di Triora, at night. C. $. D. $. E-F. Leiobunum gracile Thorell, 1876, Denmark. E. $.

F. 5 - Photographs: A-D by A.L.Schonhofer; E-F by S. Toft, all taken in the field.

MARTENS J. & SCHONHOFER A.L., The Leiobumim rupestre group

Body (Figs 1A-B, 3A-B). In both sexes dorsum smooth, minute granulation not even to be distinguished

with magnitude 250 (compare Silhavy 1981: fig. 4), few scattered pointed granules present; eye mound

in both sexes without spines or tubercles.

Dorsal pattern. Male and female (Figs 1A-B, 3A-B): contrasting black and white (see Diagnosis).

Pedipalps. Male (Fig. 4A-B): Fe with irregular rows of marked triangular spines ventrally; dorsally

nearly unarmed, ventral spines more slender than in L. gracile ; Pt slightly armed with pointed denticles

laterally; Ti markedly convex in basal part and concave at ventro-distal end, therefore forming a

well-marked S-bend, field of fine dark coloured granules all over the ventral side, well to be seen; Ta

continuously and slightly bent to ventral. Female (Fig. 4G-H): similar to male, less spines on Fe and Pt,

few spines on Ti proximally and no ventral granulation, Ta strait, slightly bent distally.

Genital morphology (Fig. 5A-C). Truncus penis stout, in ventral/dorsal view from basal opening to

insertion of glans slightly and continuously tapering; distal wings covering less than half of truncus.

Wings consist of two independent parts: a ventral shell-like plate, sharply cut off horizontally at upper

third of wing structure, from its upper margin continuously extending to lateral side of truncus, thus

partly embracing the lateral wings which extend from dorsal side of truncus via lateral side to ventral side.

Their lower two thirds are covered by the ventral plate. Surface of the upper margin of the lateral wings

and membranes adjacent to the truncus is invaginated forming a long double-walled internal sack. At

the very distal end of truncus a small membranous oval sack with a distal opening is attached ventrally.

From lateral view truncus slightly curved (concave on ventral side), slightly tapering towards glans.

Wings massively enlarged on ventral side. Glans stout, in lateral view tapering towards stylus.

Fig. 2. Distribution of the species of the Leiobunum rupestre species group. Red squares = L. rupestre

Herbst, 1799; blue circles = L. gracile Thorell, 1876; green triangles = L. apenninicum (Martens, 1969);

stippled line circumscribes area of L. subalpinum Komposch, 1998. Type localities are indicated by

abbreviated taxon names (ape = apenninicum ; gla = glabrum\ gra = gracile ; lae = laeve\ nor = norvegicum ;

rap = rupestre ; sub subalpinum ; tis = tisciae ). Arrows indicate a specified locality, if unprecisely given,

the general area is indicated ( rupestre = Saxony; gracile , laeve = Scania).

European Journal of Taxonomy 216: 1-35 (2016)

Distribution (Fig. 2)

Main area are the eastern Alps from the eastern part of Switzerland, all over Austria, the German and

Slovenian alpine parts and extending to the north-western Balkan Peninsula. Records extend southwards

to Serbia (CJM6363, CIK1336, CIK1337), Croatia (Babic 1916; Novak 2004b) and Bosnia-Hercegovina

(Martens 1978; Novak 2005). To the Northeast, the area extends into the Czech Republic: many localities

in Bohemia and Moravia, mainly in the northern and southern mountains marking the borders to Austria,

Germany and Poland (Silhavy 1981).

North of the Alps this main Alpine/Balkan area extends in isolated patches towards the northern border

of low mountain ranges in Germany (Black Forest, Schwabische Alb, Franlcenalb, Bavarian Forest and

Bohemian Forest; all Martens 1978), Fichtelgebirge (Staudt 2016), Thiiringer Wald (CJM 5392), Erz¬

gebirge (Buttner 1930), Lausitzer Gebirge (Hiebsch 1972) and northerly up to the Saxonian Elbsand-

steingebirge (CJM 4752, 4753). “Sachsen”, i.e., Saxony, is the type locality of rupestre, and the type

material of Herbst (1799) likely originated from present South Saxony.

For Poland, Stargga (1976) indicated a number of “rupestre” records in the southern mountains, the

Karlconosze (in German “Riesengebirge”) close to the Czech border. From the opposite Czech side, in

the same mountain stock, the Krkonose, Silhavy (1981) pinpointed records for (true) rupestre , which he

compared to “ glabra ” from his country. As these close-to-border Polish localities are situated in moun¬

tainous areas, too, they probably belong to rupestre as well. Besides, the Polish ‘ tisciae ’ (i.e., gracile ,

below) records are all situated in lower stretches of central and eastern parts of the country (Star^ga

1976, 2004), far from any (possible) mountainous rupestre record.

Fig. 3. Body of Leiobunum C.L. Koch, 1839, dorsal view. A-B. Leiobunum rupestre Herbst, 1799,

Germany, Mt. Arber, CJM136. A. S. B. §. — C-D. Leiobunum gracile Thorell, 1876, Denmark. C.

2 km N of Skaerbaek, CJM3530. D. $, Asp, CJM3531. — E-F. Leiobunum apenninicum (Martens,

1969), France, Alpes-Maritimes. E. <f, CJM1508. F. §, CJM2747. Drawings by K. Rehbinder.

MARTENS J. & SCHONHOFER A.L., The Leiobumim rupestre group

The L. rupestre area as depicted in Martens (1978: fig. 787) needs to be reduced, substracting the

distribution area of. L. ‘tisciae ’(fie., gracile, below), stretching from the Carpathians in a broad northerly

area to the Baltic coast up to South Finland and in the West to Denmark and northern Germany (Fig. 2).

However, the remaining distribution area of L. rupestre slightly changed with respect to the Balkan area,

then largely unknown, and new records in central Germany were added.

Ecology

This is a forest species with a high demand of air moisture. Therefore, it is confined to moist places close

to streams and creeks, on tree trunks, under rotten wood and on moist rock faces. It hardly occurs in open

areas, in agricultural land, urban gardens or on house walls in villages and towns, if not situated in moist

forests (Martens 1978), or at higher altitudes. Martens (1978) indicated height records from 260 m to

2160 m a.s.l. in the Austrian Alps, but likely included localities for the closely related L. subalpinum.

Komposch & Gruber (2004) set the Austrian uppermost rupestre limit at 1680 m a.s.l. and for sub¬

alpinum at 2100 m a.s.l.

In all cases populations of rupestre are confined to mountainous areas of different altitudes. Only in the

Saxonian Elbsandsteingebirge does rupestre live at the bottom of deeply recessed valleys which are

permanently moist and where sun radiation is largely excluded from the narrow valleys by the steep rock

faces.

Leiobunum subalpinum Komposch, 1998

Fig. 2

Leibunum subalpinum Komposch, 1998: 20.

Leiobunum rupestre Gruber, 1966: 46 (partim).

Nelima religiosa Roewer, 1910: 912 (in part: series from Tirol, GroB-Glockner).

Nelima religiosa Thaler, 1966: 79.

Leiobunum sp. - Komposch 1997: 83.

Leibunum subalpinum - Komposch 2009b: 487.

Taxonomic history

Rather uncomplicated. Because of its extreme similarity to L. rupestre , including genital morphology,

this species went unidentified in many former high-altitude reports on L. rupestre until its formal

description in 1998. Leiobunum rupestre and L. subalpinum occur in sympatry and local syntopy, but

coxal markings allow easy field identification, so no further misindications were recorded.

Diagnosis

A medium-sized Leiobunum species very similar to L. rupestre , except for black coxal markings in both

sexes and white markings on Ceph including a white area in front of the tuber oculorum in males. Coxa

IV without granules. For further details refer to Komposch (1998).

Description

Body. See Diagnosis.

Dorsal pattern. Extremely similar to L. rupestre.

Pedipalps. Very similar to L. rupestre except for a medio-basal swelling on Ta in males.

Genital morphology. Very similar to L. rupestre. The wing-free basal part of truncus shorter than in

L. rupestre.

European Journal of Taxonomy 216: 1-35 (2016)

Distribution (Fig. 2)

Restricted to the eastern Austrian Alps, mainly on the southern macroslope in the federal states of

Salzburg, Steiermark and Karnten (Komposch 1998; Komposch & Gruber 2004; Komposch 2009b)

extending into adjacent Slovenia (Novak et al. 2006). Type locality is Gofinitztal in the National Park

Hohe Tauern in Karnten.

Ecology

This a mountainous species of the eastern Alps. Its distributional range extends from about 1500 m to

2100 m a.s.l. with rare outliers down to 430 m in moist and cool ravines and up to 2200 m. Vertical

rockfaces mostly of siliceous origin are preferred (Komposch 2009a, b; Komposch & Gruber 1999).

There is a zone of sympatric and sometimes even syntopic occurrence together with L. rupestre between

500 m and 1400 m. Strongholds of L. rupestre in that area extend from 500 m to 1000 m with rare oc¬

currences below (down to 200 m) and above (up to 1400 m) (Komposch 1998).

Leiobunum gracile Thorell, 1876

Figs 2, 3C-D, 4C-D, K-J, 5D-F

Partly refers to the record within the distribution of L. gracile according to Fig. 2.

Phalangium bicolor Fabricius, 1793: 429.

Liobunum gracile Thorell, 1876: 496 (type series SMNH, examined).

Liobunum laeve Thorell, 1876: 497 (type series SMNH, examined), syn. nov.

Liobunum norvegicum Strand, 1900: 7. syn. nov.

Nelima melanogranulata Morin, 1931 (nomen nudum), 1934 (valid description) syn. to rupestre fide

Stargga 1978: 208.

Leiobunum tisciae Avram, 1968: 115 (neither declaration of type specimens nor depository for specimens,

material of the original description dedicated by Avram to J.M., now in CJM3526, examined), syn. nov.

Phalangium bicolor - Kulczynski 1876: 61. Remark: see L. rupestre.

Liobunum gracile - Tullgren 1906b: 216 (syn. with L. rupestre).

Liobunum laeve - Tullgren 1906b: 216-217 (syn. with L. rupestre).

Liobunum norvegicum - Tullgren 1906b: 217 (syn. L. rupestre).

Leiobunum tisciae - Martens 1978: 408-412 (partirn). — Star^ga 2004: 80-81. — Stol 2010b: 35. — To-

masson et al. 2014: 153.

Liobunum rupestre - Tullgren 1906a: 211; 1906b: 216. —Roewer 1910: 203-204.

Leiobunum rupestre- Roewer 1923: 890 (partirn). —Heinajoki 1944: 22-23. — Star^ga 1978: 100-103.

(partirn). — Star^ga 1979: 177-178. — Lengyel & Muranyi 2006: 121. — Enghoff 1988: 68-6 9. —

Chevrizov 1979: 14. — Spungis 2008: 21.

Nelima gracilis- Roewer 1910: 239, 250 (redescription). —Roewer 1923: 916.

Nelima laevis - Roewer 1910: 239, 251 (redescription). —Roewer 1923: 916.

Nelima norvegica - Roewer 1910: 251 (redescription). — Roewer 1923: 916-917. — Stargga 1976:

100 .

Nelima norwegica -Roewer 1910: 239. — Star^ga 1976: 100 (sic, lapsus calami).

Liobunum norwegicum - Muller 1920: 72 (lapsus calami).

Strandibunus obliquus - Bartos 1939: 309.

Nelima glabra - Kolosvary 1965: 111-113. — Kolosvary 1966a: 123. — Kolosvary & Homonnay

1967: 77-79.

Nelima nigripalpis - Kolosvary 1963: 192.—Kolosvary 1966a, b: 123.

Leiobunum glabrum - Silhavy 1981: 204-207.

MARTENS J. & SCHONHOFER A.L., The Leiobunum rupestre group

Fig. 4. Pedipalps of the Leiobunum rupestre group, whole pedipalps in lateral view, single femora in medial

view. —A-F. SS- A-B. Leiobunum rupestre Herbst, 1799, Germany, Baden-Wurttemberg, CJM1954.

A. Pedipalpus lateral. B. Femur medial. C-D. L. gracile Thorell, 1876, Denmark, Asp, CJM3531. C.

Pedipalpus lateral. D. Femur medial. — E-F. Leiobunum apenninicum (Martens, 1969), France, Alpes-

Maritimes, CJM2747. E. Pedipalpus lateral. F. Femur medial. —G-M. $ ¥• G-H. Leiobunum rupestre

Herbst, 1799, Germany, Mt. Arber, CJM136. G. Pedipalpus lateral. H. Femur medial. J-K. L. gracile

Thorell, 1876, Denmark, Asp, CJM3531. J. Pedipalpus lateral. K. Femur medial. L-M. L. apenninicum

(Martens, 1969), France, Alpes-Maritimes, CJM2747. L. Pedipalpus lateral. M. Femur medial. Arrows

indicate characters mentioned in the descriptions.

European Journal of Taxonomy 216: 1-35 (2016)

Taxonomic history

A rather complex situation developed because old species names with type localities in various areas in

Scandinavia or in (present) eastern locations were never checked in detail. This is partly due to the fact

that names were based on juvenile specimens and partly to the unavailability of former authors to access

specimens of northern (Baltic) and southern (Alpine) origin for direct comparison. Consequently,

northern populations, originally described as Liobunum gracile Thorell, 1876 and Liobunum laeve

Thorell, 1876, were permanently affiliated to L. rupestre , starting with Tullgren (1906a, b), followed by

Heinajoki (1944), Stargga (1976) and Martens (1978). The situation became even worse when Avram

(1968) described L. tisciae Avram, 1968 from the Tisza valley in Hungary. By external morphology, this

species is similar to L. rupestre , which Avram did not take into account. Martens (1978) accepted this

novelty but at that tune lacked suitable fresh material to identify populations from northern Germany and

Denmark as conspecific. Instead, he erroneously treated one species under two names, namely North

German and Scandinavian populations under L. ‘rupestre’ and those from the Carpathians and single

individuals from the British Isles under L. tiscae , besides occurrences of L. ‘rupestre’ in the Carpathian

Arc as well (Martens 1978). Silhavy (1981) was the first to recognize two very similar Leiobunum species

in (former) Czechoslovakia: L. rupestre in the West (now Czech Republic) and L. ‘glabrum’ in the East

(now Slovakia).

Thorell (1876) based his species L. gracile Thorell, 1876 and L. laeve Thorell, 1876 on juvenile

specimens from South Sweden, although he mentioned only a single male. His original material (kept at

the Museum of Natural History Stockholm) definitely concerns the types and was examined. It was

found, that it indeed only contains juveniles, as already Tullgren (1906b) assumed they represented

juveniles ofL. rupestre. Likewise, Tullgren (1906b) assumed L. norvegicum Strand, 1900 to be identical

with L. rupestre , but refrained from making a final decision, not having seen the type material. Describing

L. norvegicum , Strand (1900) depicted a female of the L. rupestre complex (from Kristiania [now Oslo];

leg. Sept. 1899) and later Martens (1978) agreed with Tullgren’s view, including L. norvegicum in the

synonymy of a broad species L. rupestre.

Considering the allopatric distribution of L. rupestre and L. ‘tisciae’ we now can firmly state that all

Leiobunum ‘rupestre’, based on material from Scandinavia, belong to L. ‘tisciae’. But as L. ‘tisciae’ is a

recently proposed name, older names are to be preferred. Albeit L. gracile and L. laeve are oldest, their

description is based on subadults (Tullgren 1906b). Yet, both have been described from Sweden, Province

of Scania, from where Tullgren (1906b) confirmed an adult specimen. Therefore, it seems a reasonable

decision to re-establish the oldest available name, Leiobunum gracile Thorell, 1876, and place L. laeve

Thorell, 1876, L. norvegicum Strand, 1910 and L. tisciae Avram, 1968 in its synonymy. L. gracile has

page and even line priority over L. laeve.

Diagnosis

A medium-sized Leiobunum species with blackish upper side, except for broad white markings of Ceph

disto-laterally in male (Fig. 3C), broad blackish irregular markings on abd area I-V of the body, in female

with mottled white markings laterally from Ceph to posterior areae, white lateral marking on abd area V

(Fig. 3D). In both sexes two paramedian small white spots on abd areae I-V, more conspicuous in male

(most similar to L. apenninicum). Contrasting yellowish underside including coxae of all appendages.

Wings of truncus penis slender, slightly tapering proximally.

Description

Body (Fig. 3C-D). In both sexes dorsal granulation consists of flat, pointed granules, regularly spaced,

well visible at 250x (compare Silhavy 1981: fig. 2).

MARTENS J. & SCHONHOFER A.L., The Leiobumim rupestre group

Dorsal pattern (Fig. 3C-D). See Diagnosis. Tu oc in both sexes smooth, without spines, in rare cases

one spine present.

Pedipalps. Male (Fig. 4C-D): Fe with irregular rows of large triangular spines ventrally often with single

hair on top; dorsally few on distal part, ventral spines broader and more massive than in L. rupestre , where

they are more slender and without hair on top. Pt with scattered denticles la and do; Ti slightly concave

ventro-distally, forming a flat S-bend, denticles and hairs on ventral side of palpal tibia light-coloured,

hard to be seen; Ta mostly straight, bent only in distal third. Female (Fig. 4J-K): in all members s im ilar

to male, but armament less conspicuous, Ti more slender, Ta strait.

Genital morphology (Fig. 5D-F). Penis similar to that of L. apenninicum. Truncus stout, from ventral/

dorsal view from basal opening to insertion of glans slightly and continuously tapering to the internal

sac of the wings; lateral wings (ventral view) markedly enlarged, smoothly rounded thus nearly egg-

shaped (irrespective the lateral membranes running down the truncus). At its very distal end the wing

structure close to the glans insertion opens to a small opening, extending into two long nearly parallel¬

sided double-walled sacks to the lower end of the wings. A lateral somewhat invaginated discharged

membrane in the distal half of the wings unites the dorso-lateral and the latero-ventral part of the wings

Fig. 5. Leiobumim , male genitalia. A-C. L. rupestre Herbst, 1799, Germany, Baden-Wtirttemberg,

CJM1954. A. Ventral view. B. Lateral view. C. Cross-section. — D-F. L. gracile Thorell, 1876,

Denmark, 2 km N of Skaerbsek, CJM3530. D. Ventral view. E. Lateral view. F. Cross-section. —

G-J. L. apenninicum (Martens, 1969), France, Alpes-Maritimes, CJM1508. G. Ventral view. H. Lateral

view. J. Cross-section. Arrows indicate the area of the respective cross-sections.

European Journal of Taxonomy 216: 1-35 (2016)

(lateral view) allowing for a possible inflation of the wing structure when the sacs are filled with secretion.

From lateral view truncus slightly curved (concave on ventral side), slightly tapering towards basis of

stylus. Wings enlarged on ventral side, slightly rounded convexly. Gians broad at truncus insertion, in

lateral view continuously tapering towards stylus.

Distribution (Fig. 2)

V _

From Slovakia (Silhavy 1981, glabra ), northern Hungary including the Tisza valley (type locality of

L. tisciae Avram; various records documented by Kolosvary 1963, 1965, 1966a, b, 1969 as nigripalpis

and glabra , and Kolosvary & Homonnay 1967), no additional records in Komposch (2004) for Hungary.

Carpathian Arc with many localities documented in collections, but few published papers available, not

very far eastward and not in the southern lowlands (Muranyi & Lengyel 2006; Weiss 1996; Cirdei 1960).

In the Southwest the area extends into Serbia (Bor Dubasnica, Mala tisnica, CJM 6363). There are

scattered records for the western part of Ukraine, mainly as an extension of the Carpathian area (Bartos

1939; Cirdei 1960; Star^ga 1978; Morin 1931, 1934; L. Koch 1870). The many records in Poland pertain

to the eastern two thirds of the country, extending to the Baltic Sea (Stargga 1978, 2004, rupestre; for

true Polish rupestre see above) and including former WestpreuBen. Occurrences in Russia are reported

from former OstpreuBen (Le Roi 1914) and from St. Petersburg (CJM 3032), extending to Northeast

Poland (Star^ga 1976,2004). Estonia: records along the Baltic Sea Coast (Tomasson etal. 2014). Latvia:

partly detailed locality maps (Tumss 1963; Spungis 2008). Finland: only in the southern part, with

detailed map based on monitoring records (Heinajoki 1944), presence confirmed by Uddstrom et al.

(2013). Germany: records along the Baltic Sea Coast and its hinterland (up to Hamburg), dating back to

Kraepelin (1896), Le Roi (1914) and Rabeler (1929), but apparently presently very rare, last record in

Schleswig-Holstein, Liibeck in 2007 (CJM 6111; see Martens 1978). Denmark: early records by Hansen

(1884), more recent records from all over the country remained largely unchanged between about 1960

(Meinertz 1964) and 1987 (Enghoff 1988), with slight distributional differences, formerly missing from

Funen and rare in South Jutland and Himmerland, but very co mm on and abundant there in 1987 (Enghoff

1988), including Funen, South Jutland and Himmerland. Since then, it has vanished, has not been

recorded for many years, but it was rediscovered recently and is presently extremely rare. In 2008, four

individuals were present on four out of 64 sites (Enghoff et al. 2014). Sweden: except for the early

records of Thorell (1876) in Vastergotland, Ostergotland, Skane and Smaland, there is a single new one

from Uppsala in 2009 (examined, H. Enghoff leg.). Norway: Stavanger (SMF) and Oslo (= Kristiania;

Strand 1900, norvegicum). Great Britain: Derbyshire (Martens 1978), unrecorded until 2008 when it

was found around Aberdeen and Dunbennan, Huntly (Davidson 2009, CJM 6358-6361), in Scotland

apparently spreading and in Great Britain presently known from seven counties (British Arachnological

Society 2016).

The present distribution of L. gracile and that in the recent past obviously represent two different

ecological-defined origins. One belongs from lowland to montane and riverine forests, the other one

comprises secondary urban habitats to which the species was (most likely) transferred by human activity.

Such “anthropogenic” populations may suddenly increase, occupy a large area and sometimes collapse

again, for reasons little understood. Others are spot-like with little influence on local faunas. At least for

a period of roughly 120 years such dynamics can tentatively be reconstructed (see Discussion).

Ecology

Aeuryoecious species. In Slovakia, including the Carpathian Arc, recorded from sheltered places in old-

growth forests on tree trunks and on rock faces (Silhavy 1981). It is co mm on in the inundation zone of

rivers like the Tisza, e.g., frequently reported from tree hollows of Populus (Avram 1968, tisciae ) and

Salix trunks (Avram 1968; Csizmazia et al. 1966, nigripalpis ; Kolosvary 1963, 1965, 1966a, b). This

also holds true for East Poland where gracile lives in forested habitats (Neple, Bug River, AXLS 958).

Weiss (1996) indicated its occurrence from Romanian Transylvania forests of the hilly and montane belt.

MARTENS J. & SCHONHOFER A.L., The Leiobumim rupestre group

In all other areas, which we believe to be secondary, L. gracile is confined to strongly modified and

human-influenced habitats. There it competes with other species preferring house walls like Opilio

saxatilis C.L. Koch, 1839, O. parietinus (Degeer, 1778) and recently with O. canestrinii Thorell, 1876.

In central Europe the latter presently is by far the dominant species in human habitats and may have

partly caused the noticeable decline of O. saxatilis , O. parietinus and L. gracile (see Discussion).

Leiobunum apenninicum (Martens, 1969)

Figs 1C-D, 2, 3E-F, 4E-F, M-L, 5G-J

Nelima apenninica Martens, 1969: 409 (type series SMF, examined).

Leiobunum rupestre - Trossarelli 1943: 54. — Gozo 1908: 135.

Nelima apenninica - Martens 1978: 424, figs 803-804, 809. — Chemini 1980: 79 (description of

female). — Komposch 2009: 457.

Taxonomic history

Though only recently described, the status of this large species was accepted by subsequent authors

(Chemini 1980; Komposch 2009). However, similarity of genital morphology with L. ‘ tisciae 7 (now

L. gracile , Martens 1978) later led Martens to the assumption that the scattered distribution of apenninica

might represent disjunct area parts of L. gracile. This remained unpublished due to the emerging complex

taxonomic situation in L. gracile/L. rupestre. Yet, many details separate both species as is further

confirmed by molecular genetic analysis (J. Martens, unpublished data).

The generic affiliation of apenninica is difficult to establish. Certainly, it does not belong to the European

bulk of small-bodied Nelima species like N. sylvatica Simon, 1879, N. gothica Lohmander, 1945 and

N. sempronii Szalay, 1951, but is a large and long-legged species similar to L. gracile , less to L. rupestre.

Due to the lack of coxal denticles in the type material and according to the present definition of

sclerosomatid genera it was originally placed in Nelima. However, based on only a single external

character, this is an artificial affiliation, that we find especially compromised in the species-pair gracile

and apenninica. Leiobunum gracile shows a strong variation of number and placement of coxal denticles

down to zero (CJM6357, female) and their frequent appearance in Nelima apenninica (AXES 1739 two

to seven on coxa IV, in both sexes) underlines the false generic placement of apenninica. Consequently,

in accordance with similar genital morphology and external characters we prefer to place apenninica in

Leiobunum.

Diagnosis

A medium-sized Leiobunum species with blackish upper side, except for broad white markings of Ceph

disto-laterally in male (Figs 1C, 3E), broad blackish irregular markings on abd areae I-V of the body. In

female with mottled white markings laterally from Ceph to posterior areae, white lateral marking on abd

area V and a white cross stripe all over areae VI (Figs ID, 3F). In both sexes a series of para-median

small white spots on the abd areae I-V, more conspicuous in male (a character similar to L. gracile).

Contrasting yellowish under side including coxae of all appendages. Wings of truncus penis robust,

slightly tapering proximally.

Description

Body (Figs 1C-D, 3E-F). In both sexes dorsal granulation consisting of flat, pointed granules, regularly

spaced, well to be seen with 250x (comparable to L. gracile , see Silhavy 1981: fig. 2); eye mound in

both sexes with 1-3 small spines on each side, rarely absent.

Dorsal pattern (Figs 1C-D, 3E-F). See Diagnosis.

European Journal of Taxonomy 216: 1-35 (2016)

Pedipalps. Male (Fig. 4E-F): Fe with few massive spines, normally restricted to third ventro-distal

part, only single spines on dorso-distal side or absent; Pt unarmed except for few small spines dorsally;

Ti slighty inflated, rather short thus appearing pretty massive, brush-like hair field on ventral side,

a peculiar character of its own; no spines clearly visible, likely for being of the same light colour as

the embedding hair field; Ta continuously bent ventrally over its whole length. Female (Fig. 4M-L):

Similar to male. Fe nearly smooth, spines even less than in male, tibia less inflated and slightly concave

ventrally, no hair field; tarsus straight, distally inconspicuously bent ventrally, more slender and slightly

longer than in male.

Genital morphology (Fig. 5G-J). Penis similar to L. gracile. Truncus stout and parallel-sided in the

basal half from where slightly and continuously tapering to the lateral lowest parts of the wings, from

there slightly enlarged to middle of the wings, then tapering to basis of glans. Lateral wings (ventral

view) markedly enlarged, nearly parallel-sided over nearly their total length, pretty sharply merging to

distal part of truncus. At the disto-lateral margin of the wings (close to the truncus/glans joint) wings

open to a narrow slit leading to two ventro-lateral pockets. A lateral membrane in the distal half of

the wings unites the dorso-lateral und the ventro-lateral part of the wings which allows for a possible

inflation of the wing structure when the pockets are filled with secretion. From lateral view truncus

slightly curved (concave on ventral side), slightly tapering towards middle part, then enlarging again.

Wings massively enlarged on ventral side, slightly rounded convexly. Gians slender, in lateral view

tapering towards stylus.

Distribution (Fig. 2)

It was long considered a rare and local species (Martens 1969; Komposch & Gruber 2004; Komposch

2009; Chemini 1980) and only a few specimens were available for the original description from the

Apennines (Martens 1969). Even fewer became known from the apparently isolated area at the Italian

border to Austria (Chemini 1980) and in Austria itself (Martens 1978; Komposch 2009) and the

Southwest Alps (Martens 1978). Presently, the area seems to be subdivided into three major parts, viz.,

French and Italian southwestern Alps, southern Austrian (eastern Tyrol) and Italian Alps (South Tyrol),

and scattered records in the Apennines chain. Type locality is Pratovecchio in the Apennines. Yet, there

was little additional locality information since its original description until the species was recently

rediscovered in the Southwest French Alps in high frequency, namely in the Mercantour National Park

and the Vallee des Merveilles (CJM; ALS leg., JM leg). Disregarding its specific habitats and general

low sampling activity for long-legged species in the Northwest Alps, the species might have been

neglected in large parts of its distributional area. Prior to its description, L. apenninicum has likely been

confused with L. rupestre , as is obvious from Trossarelli’s (1943) records for the southwest Alps. The

isolated record from Calabria (Gozo 1908, L. rupestre ; included in Fig. 2) requires confirmation.

Ecology

This is a forest species. During the day males rest on tree trunks and rock faces; females, though not

exclusively, prefer to hide on the ground in leaf litter and thus are not easy to locate. Consequently, only

males have long been known, because they are easier to spot and collect. The species is locally pretty

common wherever the daytime microhabitats of females are found. Otherwise, it is readily encountered at

night, and this technique revealed large populations in the Mercantour National Park in the French Alps.

Discussion

Systematics of the four central European species Leiobunum rupestre , L. subalpinum , L. gracile and

L. apenninicum highlights general problems related to European sclerosomatid taxonomy. Firstly, in this

case old, but available names, have been largely neglected and were not compared to species known from

the area. Subsequent species proposals, i.e., Leiobunum norvegicum or Leiobunum tisciae ignored this

MARTENS J. & SCHONHOFER A.L., The Leiobunum rupestre group

point and caused considerable taxonomic confusion. Old names, though available, were not considered in

detail and simply put aside. On the other hand, Leiobunum subalpinum , a species very similar to L. rupestre

but not rare at all in its limited alpine area, escaped taxonomic recognition for about 200 years.

Furthermore, the artificial division of Leiobunum into Leiobunum s. str. and Nelima proposed by Roewer

(1910) and maintained for a century seems to be inappropriate in the case presented here. The Roewerian

separation is based solely on the presence (Leiobunum) or absence (Nelima) of coxal denticles, but does

not place “ Leiobunum” and “ Nelima ” species into reasonable morphological and much less preferably

monophyletic units, which is substantiated by results of modern molecular genetics (Hedin et al. 2012).

In our example, “ Nelima ” apenninica normally does not generally bear such denticles, but its male

genital morphology is astonishingly similar to that of L. gracile to an extent that one might even suppose

conspecifity. “ Leiobunum ” gracile is reported to have highly variable numbers of coxal denticles (Avram

1968) and partly none, while our material also revealed several individuals of “Nelima” apenninica to

have few denticles, too. Molecular genetic analyses indeed demonstrate close relations (J. Martens,

unpublished data), confirming the specific differences. The taxon apenninica belongs to Nelima only by

Roewer’s genus definition, genetically and genital-morphologically it is affiliated to a group of central

European Leiobunum. However, “LeiobunumT , presently a holarctically distributed genus, is

heterogeneous as well and refers to a number of morphologically and genetically distinct lineages,

which are far from being disentangled (Hedin et al. 2012). From our material it is also apparent that an

increase of the number of denticles, as well as their presence on more than one coxa is linked to an

increased body size of the individual, at least in apenninica and gracile. Such a variable character is

certainly inappropriate to serve for differentiation of higher level taxa within a highly heterogeneous

phylogenetic group. A molecular-based re-organization, not only of the European Leiobunum/Nelima

complex, is in urgent need.

The distributional analysis revealed that Leiobunum gracile and L. apenninicum , though closely related,

show a very different distributional pattern. Leiobunum apenninicum is a conservative forest species,

apparently confined to a few scattered areas and presently with little perceivable tendencies to enlarge its

distribution.

The closely related L. gracile , however, massively enlarged its area in close affinity to man-made habitats

around the Baltic Sea coast. In this area of expansion its populations considerably fluctuate in numbers

and it can constrict its area down to local disappearance. Parts of the Carpathian Arc and the adjacent

mountains in Southwest Poland and Slovakia and low-altitude riverine forests in Hungary may represent

the original area of the species, though this is difficult to substantiate. “Natural” populations may be

indicated by hilly and montane forests, also riverine localities where high population densities are

obvious (Avram 1968; Csizmazia & Kolosvary 1966; Kolosvary 1963, 1965, 1966a, b, 1969; Kolosvary

& Homonnay 1967).

However, it remains arbitrary to solidify area limits of an original and an expansive population based on

ecological comparison of historical records. It is thus only hypothesized that starting from the Carpathians

the Baltic coast area and its hinterland have been colonized, possibly by help of human-mediated

transport. All Baltic authors (Rabeler 1929; Kraepelin 1896; Ee Roi 1914; Heinajoki 1944; Tumss 1963;

Meinertz 1964; Enghoff 1988; Enghoff et al. 2014; Stargga 1978, 2004) emphasize the close proximity

to human settlements in their area, such as in gardens, specifically on house walls and similar

environments, while forest habitats played a minor role in all ecological characterisations. In the Baltic

area, this presumed invasion of new areas was successful, but to the West, the North Sea coast was not

reached, at least not in a continuous belt, with the westernmost localities around Hamburg (Kraepelin

1896). Isolated outposts are known from the British Isles (Martens 1978) including Scotland (British

Arachnological Society 2016), where the population seems to prosper.

European Journal of Taxonomy 216: 1-35 (2016)

A reasonable proof of newly gained territories is the fact that in the newly colonized areas population

densities are high in “disturbed” habitats, remain stable for a certain period and after years or even

decades collapse or finally completely disappear. This holds true for the population in Denmark, stable

at least from the mid 1960s (Meinertz 1964)to 1987, when Enghoff (1988) undertook another monitoring

and found an even more extended territory inhabited. Presently, L. gracile has largely disappeared from

De nm ark (Enghoff et al. 2014). Likewise in northern Germany only a few isolated records were reported

between 2000 and 2009 (Staudt 2016), while the species appeared common at the beginning of the last

century (Kraepelin 1896; Rabeler 1929; Le Roi 1914). Other occurrences remained spot-like from the

very beginning and apparently disappeared soon after the first discovery in Great Britain (Martens 1978,

tisciae) or gained new area spots far away in Scotland and other parts of the U.K. (British Arachnological

Society 2016).

Reasons for such extreme population fluctuations are unclear and have not been surveyed in detail.

Population decline was obvious for long prior to the invasion of Opilio canestrinii to central Europe,

which started about 1970 (Martens 1978, as ravennae, first German records in Stuttgart 1974 CJM1619;

Mainz 1975 CJM1525; Berlin 1976 CJM1553). For the year 1987 Enghoff (1988) stated that L. gracile

and O. canestrinii lived side by side in high population densities in Denmark. Only gracile disappeared,

canestrinii remained to the present and is common everywhere (Toft 2004, pers. comm.). However, it

must be taken into account that long-lasting effects of high population density of O. canestrinii may

have caused the decline of L. gracile. The same is assumed for Opilio parietinus in central Europe.

The temporal invasion of the Baltic dates back for at least 120 years, probably considerably longer.

Kraepelin (1896) mentioned gracile as not being rare for the Hamburg area whereas Heinajoki (1944)

stated that gracile invaded Finland and gained northward territory only in the 1920s. Monitoring of the

Finnish arachnid fauna was pretty detailed at that tune and earlier occurrences would have been certainly

noticed as indicated by Heinajoki (1944), who documented the extension to about 61.6° N up to

Hameenkyro, until today the northe rnm ost known record (Uddstrom et al. 2013). For Denmark records

date back to the 19 th century, when Hansen (1884) mentioned the species.

Acknowledgements

We are much indebted to A. Ausobsky, A. Babalean, H. Enghoff, M.B. Davidson, S. Huber, P. Jager,

I. Karaman, Ch. Komposch, M. Lemke, H. Pieper, W. Star^ga and S. Toft who provided material and

information for the study of this difficult group. T. and S. Novak, R. Ozimec, J. Wolff, A. Grabolle,

S. Huber, M. Isaia and M. Paschetta further helped with field collections. H. Enghoff translated Danish

text and provided data at an early stage of manuscript work. M. Grasshoff and P. Jager allowed access

of the arachnid collection of the Senckenberg Natural History Museum Frankfurt. K.A. Johanson and

G. Lindberg at the Natural History Museum Stockholm gave advice and access to Thorell's types.

K. Rehbinder created the beautiful drawings of dorsal body patterns. Wagner Foundation and Feldbausch

Foundation at Fachbereich Biologie of Mainz University granted annual financial support for ecological

and systematic studies on harvestmen in Europe and Asia to J.M. A.L.S. collected European Leiobunum

in the Alps with the help of traveling funds from DAAD (2006) and EDIT/ATBI+M (2008). Finally, we

greatly acknowledge the helpful remarks and hints of two anonymous reviewers. We heartily thank all

friends, colleagues and institutions.

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Manuscript received: 2 February 2016

Manuscript accepted: 5 April 2016

Published on: 27 July 2016

Topic editor: Rudy Jocque

Desk editors: Niko Korenhof & Kristiaan Hoedemakers

Printed versions of all papers are also deposited in the libraries of the institutes that are members of the

EJT consortium: Museum national d’Histoire naturelle, Paris, France; Botanic Garden Meise, Belgium;

Royal Museum for Central Africa, Tervuren, Belgium; Natural History Museum, London, United

Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium; Natural History Museum of

Demnark, Copenhagen, Demnark; Naturalis Biodiversity Center, Leiden, the Netherlands.

European Journal of Taxonomy 216: 1-35 (2016)

Appendix 1 - List of material and distributional data

Remarks

The list incorporates investigated material and references, which we judged sufficiently reliable. Old

or alternative locality names, interpretations and accuracy of coordinate-reconstruction are indicated in

parenthesis. Important taxonomic material, such as types, are indicated in bold.

Acronyms of institutions cited in the list

AXLS = Coll. A. Schonhofer, Ma in z. Germany

BGZM = Biozentrum Grindel and Zoologisches Museum, Universitat Hamburg, Germany

CIK = Coll. Ivo Karaman, Novi Sad, Serbia

CJM = Coll. Jochen Martens, Mainz, Germany

MBN = Museum Bergen, Norway

MFSNU = Museo Friulano di Storia Naturale, Udine, Italy

MNHN = Museum national d’histoire Naturelle, Paris, France

MTSN = Museo Tridentino di Scienze Naturali, Trento, Italy

MNV = Museo Civico di Storia Naturale di Verona, Italy

SMF = Senckenberg Forschungsinstitut Frankfurt a. Main, Germany

UNFLAT = Department of Zoology and Animal Ecology, Faculty of Biology, University of Latvia

ZMB = Museum fur Naturkunde Berlin, Germany

Leiobunum apenninicum (Martens, 1969)

AUSTRIA: Osttirok Arnbach near Sillian, 1100 m, 46.74° N, 12.39° E, leg. A. Ausobsky, 15 Jun. 19??

(Martens 1978); Defereggental, banks of river Schwarzach below Mariahilf, 1670 m, 46.913° N, 12.31° E,

leg. 31 Aug. and 1 Sep. 2007 (Komposch 2009).

FRANCE: 5 33 , 4 §§, 2 juv., Pep. Isere . Le-Sappey-en-Chartreuse, direction to Sarquenas, 1158 m,

mixed coniferous and Fagus forest, under bark and logs, 45.273° N, 5.766° E, leg. 31 Aug. 2008

(AXLS30); 3 juv., Pep. Alpes-Maritimes. buffer area of the Mercantour National Park, focal site La

Miniere de Valaura (9), 1476 m, mixed forest on north facing slope, under stones and logs, 44.069° N,

7.132° E, leg. 5 Sep. 2008 (MNHN); 4 juv., same locality, below Col de Turini, 1513 m, coniferous

forest, under logs and bark, 43.977° N, 7.386° E, leg. 3 Sep. 2008 (AXLS85); 1 3 , Foret de Pei'ra Cava,

1500 m, 43.93° N, 7.37° E, leg. 18 Aug. 1972 (CJM1508; Martens 1978); 2 33 , 1 $, Col de Turini,

1500 m, Picea forest, below pass, 43.978° N, 7.390° E, leg. 19 Sep. 1982 (CJM2747); 28 33 , 2

15 juv., Mercantour National Park, focal site Valmasque (7), 1905 m, NW exposed slope, very humid,

indicated by predominance of Alnus viridis , at stems of Larix , 44.111° N, 7.473° E, leg. 6 Sep. 2008

(CJM6295).

ITALY: 6 33 , Prov. Arezzo. Casentmo . Bibbiena vicinity, P. di Mandioli (interpreted as Passo di

Mandrioli), 1050 m, rivulet in Fagus forest, 43.78° N, 12.01° E, leg. 18 Oct. 1975 (CJM3287); 2 33 ,

Central Apennin . Matese Mts., Roccamandolfi, 820 m, 41.50° N, 14.35° E, leg. 27 Jun. 1967 (MV;

Marcellino 1971; Martens 1978); 1 3 , Sotto Rifugio Campitello, 1250 m, 41.463° N, 14.394° E, leg. 22

Sep. 1967 (MV; Marcellino 1971; Martens 1978); Emilia Romagna . Prov. Bologna, 1 $, “Corno alle

Scale” Park (coordinates inaccurate), 44.14° N, 10.83° E, leg. 2005 (online source Tree of Life,

photographic evidence ID11332); 1 3 , Forli-Cesena, Parco Nazionale Foreste Casentinesi, Poggio Scali,

Monte Falterone, 1520 m, 43.845° N, 11.79° E, leg. Sept. 1963 (SMF16433; Martens 1969; cf. Kirchner

& Salzer 1967; holotype of Leiobunum apenninicum (Martens, 1969)); 2 33 , same locality,

Campigna, 1050 m, 43.87° N, 11.73° E, leg. Sept. 1963 (SMF16434; Martens 1969, paratypes); 1 3 ,

1 $, Pratomagno, E Firenze, Vallombrosa (interpreted as the abbey), 1100 m, Abies forest, 43.732° N,

MARTENS J. & SCHONHOFER A.L., The Leiobunum rupestre group

11.558° E, leg. 17 Oct. 1975 (CJM3282); Prov. Imperia . 9 ad., 1 juv., Monesi di Triora, 1300 m, stream

valley before village, night catch, 44.081° N, 7.761° E, leg. 30 Aug. 2014 (AXLS1733); 1 3 , first stream

valley before village, 1400 m, in deep stony gravel and sieved from litter in-between, 44.071° N,

7.748° E, leg. 30-31 Aug. 2014 (AXLS1701); 20 33 , 31 $$, 3 juv., last valley before Monesi di Triora,

1287 m, sitting on dry leaf litter (often under fallen twigs) in mixed forest, 44.081° N, 7.767° E, leg. 9

Sep. 2008 (CJM6308); 1 3 , Monesi di Triora, 1304 m, deciduous forest ( Fagus j, under stones, 44.082°

N, 7.771° E, leg. 8-9 Sep. 2008 (AXLS111); 1 $, c/o Monesi, 1320 m, 7.761° N, 44.081° E, leg. 24 Jun.

2011 (AXLS1150); 1 3 , Alpi Cozie, Crissolo, Monviso, 1300 m, 44.67° N, 7.10° E, leg. July 1967 (MV;

Marcellino 1975); 1 3, N Pinerolo, Giaveno, W Forno (interpreted as Forno), 950 m, 45.04° N, 7.24°

E, leg. 9 Oct. 1972 (CJM1576; Martens 1978); 8 33 , 4 5$, 4 juv., Alpi Marittimi. National Park Alpi

Marittime, focal site Palanfre, 1501 m, old N-exposed Fagus sylvatica forest, sitting on leaf litter,

44.195° N, 7.494° E, leg. 10 Sep. 2008 (MNHN); 1 juv., focal site Vallone d’ Argentera, 1580 m, coniferous

forest, under stones, 44.185° N, 7.269° E, leg. 12 Sep. 2008 (MNHN); 2 33 , 2 §?, Nava, 888 m, night

catch alongside deciduous forest and stream, mostly on road, 44.099° N, 7.874° E, leg. 7 Sep. 2008

(AXLS108); 4 33 , 2 $$, Val de Valasco, Rifugio Valasco, 1800 m, N-facing slope with large boulders,

Alnus viridis and Myrtillus shrubs, under stones and in crevices, 44.196° N, 7.233° E, leg. 28 Aug. 2014

(AXLS1739); 3 33 , Trentino-SudtiroL betw. Nigerpass and Karerpass, E Welschnofen, 1720 m, 46.43°

N, 11.58° E, leg. 28 Aug. 2005 (CJM4670); 1 3 , 1 ?, surroundings of Palu del Fersina, 1300-1400 m,

46.13° N, 11.34° E, leg. 8 Sep. 1973 (MTSN, Chemini 1980); 1 3 , surroundings of Paneveggio, 1500 m,

46.31° N, 11.70° E, leg. 28 Aug. 1973 (MTSN, Chemini 1980). Calabria . Valletta della grotta della Melia,

38.23° N, 15.76° E (Cantoni 1882; Gozo 1908; orig. det. as L. rupestre and is here affiliated to L.

apenninicum ; requires confirmation).

Leiobunum gracile Thorell, 1876

Remarks

Leiobunum gracile was mostly considered under the names L. rupestre and L. tisciae. These names we

used for the original identification of records in the literature. The repetitive information was omitted,

while more unusual species taxa names and identifications ( glabra , nigripalpe , etc.) are given. If neces¬

sary, please refer to the respective publication.

DENMARK: 3 33 , 2 $ $, 1 juv., 2 kmN of Skaerbaek, coniferous forest, 55.17° N, 8.76° E, leg. 16-30 Sep.

1987 (CJM3530); 1 3 , 3 ??,Asp, indoors, 56.824°N, 9.936°E, leg. 19 Sep.-6 Oct. 1987 (CJM3531);

Bornholm, 55.16° N, 14.87° E (Meinertz 1964); 4 §§, Givskud, 55.82° N, 9.37° E, leg. 16 Sep. 1987

(CJM6232); 2 33 , 3 ??, Heming, 56.13° N, 8.97° E, leg. 16 Oct. 1987 (CJM6230); 1 3 , 2 ??,

Copenhagen (coordinates inaccurate), 55.66° N, 12.59° E (ZMB12642); 2 33 , 1 ?, Copenhagen, leg.

10 Sep. 1984 (CJM4799); 14 ad, Stjse, 56.12° N, 8.97° E, leg. 30 Sep. 1987 (CJM6231); 1 3 , Torvekj<er

v. Klitmoller, 57.03° N, 8.55° E, leg. 2 Dec. 2006 (CJM6229).

ESTONIA: All Estonian records refer to Tomasson et al. 2014 based on 3-4 sampling periods of 4-7

months in 2008-2011. 3 ex., Harju, Uksnurme, unmanaged old apple orchard next to cultivated grass¬

land, 59.295° N, 24.628° E; 15 ex., Hiiu, Kerema, moist alvar grassland with juniper brush, 58.891° N,

22.948° E; 116 ex., Jogeva, Pataste, moist forest dominated by alder, birch, bird cherry, spruce, 58.581° N,

26.778° E; 18 ex., Laane, Kunila, Moist broad-leaved forest, 58.631° N, 23.811° E; 1 ex., Parnu, Nigula

NR, paludified pine forest with Polytrichum , 58.007° N, 24.712° E; 11 ex., Parnu, Tostamaa, next to

brush covered ditch bank surr. by fallow grassland, 58.336° N, 24.006° E; 56 ex., Saare, Igakiila, Pine

forest with Corylus, 58.601° N, 23.128° E; 18 ex., Saare, Orissaare, patch of deciduous managed forest

in an urban area, 58.555° N, 23.087° E; 1 ex., Saare, Viidumae, edge of a spring fen, 58.294° N,

23.089° E; 108 ex., Tallin, 59.44° N, 24.75° E (Star^ga 1978); 108 ex., Tartu, Maiorg, spruce and aspen

dominated old forest along a rivulet valley, 58.278° N, 26.334° E; 136 ex., Tartu, Sudaste, Oxalis-typQ

European Journal of Taxonomy 216 : 1-35 ( 2016 )

drained spruce forest, 58.329° N, 26.940° E; 11 ex., Valga, Puka, edge of paludified forest, Filipendula

type, 58.041° N, 26.207° E; 48 ex., Voru, Leoski, edge of dry boreal forest, Vaccinum myrtillus type,

57.709° N, 27.139° E.

FINLAND: 2 juv., Espoo, 60.21° N, 24.65° E, leg. 17-18 Jul. 1940 (Heinajoki 1944); Hame . Forssa,

60.82° N, 23.63° E (Hippa 1975); Tammela, 60.80° N, 23.77° E (Hippa 1975); Ypaja, 60.80° N, 23.28° E

(Hippa 1975); Hameenkvro . 1 $, 61.64° N, 23.20° E, leg. 1943 (Heinajoki 1944); Koiiarvi. 60.94° N,

23.66°E (Hippa 1975);N. Helsinki .30 150 $$, 173 juv.,Elaintarhanhuvila 6, 60.18°N,24.94°E, leg.

1935-40 (Heinajoki 1944); 1 juv., Kapyla, indoors at wall, 60.21° N, 24.95° E, leg. 3 Aug. 1925 (Heinajoki

1944); Oulunkyla and Pakinkyla, on path and indoors, 60.23° N, 24.94° E, leg. late summer 1943 (Heinajoki

1944); 1 S, Ullanlinna, indoors at wall, 60.16° N, 24.95°E, leg. 25 Sep. 1926 (Heinajoki 1944); Southern

Finland . Humppila, 60.93° N, 23.37° E (Hippa 1975); Jokioinen, 60.80° N, 23.48° E (Hippa 1975); St

Hameenkvro . 1 ad., on a path, 61.635° N, 23.193° E, leg. 20 Sep. 1943 (Heinajoki 1944); Turku (coordinates

inaccurate), 1 S, 1 $, 60.45° N, 22.26° E, leg. 25 Sep. 1926, 9 May 1927 (Heinajoki 1944); Western

Finland . Somero, 60.63° N, 23.52° E (Hippa 1975); Urjala, 61.08° N, 23.54° E (Hippa 1975).

GERMANY: Hamburg (three localities), large numbers (coordinates inaccurate), 53.56° N, 9.87° E

(BGZM); Barmbeck, 53.58° N, 10.03° E (Kraepelin 1896); Bahrenfeld, 1 Stadtteil von Hamburg,

53.57° N, 9.91° E (SMF-R.1255); St. Georger Kirchhof, 53.557° N, 10.007° E (Kraepelin 1896);

Blankenese, 53.56° N, 9.81° E (Kraepelin 1896); Mecklenburg-Vorpommern . 1 ex., Rostock, house

garden in nesting box (Rabeler 1929); 4 ex., Rostock, old cemetery at Saarplatz (now abandoned), at

houses and walls, 54.085° N, 12.117° E, leg. 8 Oct. 1927-1929 (Rabeler 1929); 1 ex, Rostock, yard of

the Entomology Seminar, 54.088° N, 12.134° E, leg. 7 Oct. 1927-1929 (Rabeler 1929); 1 ex., Rostock,

indoors, leg. 7 Nov. 1927-1929 (Rabeler 1929); 3 ex., Rostock, Wallanlagen, in cracks of wall, 54.087°

N, 12.132° E, leg. 13 Sep. 1927-1929 (Rabeler 1929); Wismar, 53.89° N, 11.45° E (Le Roi 1914);

Schleswig-Holstein . Bad Oldesloe, 54.10° N, 10.79° E (BGZM; Kraepelin 1896); 1 $, 2 $$, Kiel,

Institute ofZoology, 54.33° N, 10.1° E, leg. 1 Sep. 1964 (CJM949, CJM6172); l l $, Ltibeck, former

School of Navigation, on wall, 53.861° N, 10.689° E, leg. 9 Oct. 2007 (CJM6111); 1 S, 4 $ $, Neumunster

(72), 20 m, 54.07° N, 9.99° E, leg. 17 + 27 Sep. 1964 (CJM6171); Neumunster(?) (coordinates inaccurate),

1 (?, 54.07° N, 9.99° E, leg. Sep. 1967 (CJM948).

GREAT BRITAIN: Scotland . Aberdeenshire . 2 SS, 1 ?, Aberdeen, Botanic garden, 57.16° N, 2.09° W,

leg. 20 Sep. 2008 (CJM6359); 2 $$, same data (CJM6358); 4 33, 1 ¥, same data (CJM6357); 2 <$<$,

1 5, Aberdeen, Torry, Greyhope House, 57.142° N, 2.062° W, leg. 9 Oct. 2008 (CJM6361); 1 (?, 3 $

same data, leg. 4 Oct. 2008 (CJM6360); 1 Dunbennan, Huntly, Cemetery, 57.45° N, 2.79° W, leg.

12 Sep. 2008 (CJM6362); Derbyshire . 13 ex., between Leek and Stoke on Trent (coordinates inaccurate),

53.06° N, 2.12° W, leg. Sep. 1971 (CJM1739) (Martens 1978).

HUNGARY: All Hungarian records of Kolosvary (1963, 1966a, b), Kolosvary & Homonnay (1967) and

Csizmazia etal. (1966) are originally det. as Nelima nigripalpe , of Kolosvary (1965) orig. det. as Nelima

glabra and of Kolosvary (1969) orig. det. as Nelima glabra and TV. nigripalpe considered as syn. Northern

Hungary : Biikk Mts., Bankut, Veres sar berc, 48.09° N, 20.49° E, leg. 2 Nov. 2005 (Lengyel & Muranyi

2006, orig. det. as L. rupestre); Tisza River and surroundings. Kislcore, 47.49° N, 20.51° E, leg. Aug.

1968 (Kolosvary 1969); 1 ex., Komoro, inundation zone, Salix forest, in humus, 48.31° N, 22.09° E, leg.

18-31 Jul. 1965 (Kolosvary 1965); Nagyrev, 46.94° N, 20.15° E, leg. Aug. 1968 (Kolosvary 1969);

river kilometre 507, „Koldus“ forest near Tiszadob, inundation zone, tree hollows of Salix stem,

48.03° N, 21.18° E, leg. 18-19+23+31 Jul. 1964 (Csizmazia et al. 1966); many ex. river kilometre

601-602 (near Dombrad), inundation zone, tree hollows of Salix , 48.24° N, 21.92° E, leg. 18-31 Jul.

1965 (Kolosvary 1965); river kilometre 602-604 (1-3 km NE Dombrad), inundation zone, at walls of

cottage, 48.26° N, 21.94° E, leg. 18-31 Jul. 1965 (Kolosvary 1965); 1 ex. river kilometre 620, Tuzser,

inundation zone, moulting at branch of Salix, 48.34° N, 22.11° E, leg. 18-31 Jul. 1965 (Kolosvary

MARTENS J. & SCHONHOFER A.L., The Leiobunum rupestre group

1965); 1 ex., Szabolcsveresmart, inundation zone, Quercus forest, under log, 48.31° N, 22.02° E, leg.

18-31 Jul. 1965 (Kolosvary 1965); Tiszabercel, 48.17° N, 21.66° E, leg. Aug. 1968 (Kolosvary 1969);

Tiszacsermely, 48.23° N, 21.81° E, leg. Aug. 1968 (Kolosvary 1969); Tiszadada, inundation zone, listed

as terricole species, 48.04° N, 21.24° E, leg. 1963-1964 (Kolosvary 1966b); 1 ex., Tiszadob, inundation

zone, listed as terricole species, 48.03° N, 21.18° E (Kolosvary 1963); Tiszafoldvar, 46.99° N, 20.22° E,

leg. Aug. 1968 (Kolosvary 1969); Tiszahat, Endes-Puszta, on Rubus leaves, 48.29° N, 22.23° E, leg.

1 Aug. 1966 (Kolosvary & Homonnay 1967); Tiszahat, Gergelyiugornya, river kilometre 698, in forest,

48.13° N, 22.34° E, leg. 27 Jul. 1966 (Kolosvary & Homonnay 1967); Tiszahat, Gergelyiugornya, river

kilometre 698, in forest, skins in tree hollows of Salix and Quercus , 48.13° N, 22.34° E, leg. 27 Jul. 1966

(Kolosvary & Homonnay 1967); Tiszahat, Lonya, inundation zone, 48.32° N, 22.25° E, leg. 21 Jul. 1966

(Kolosvary & Homonnay 1967); Tiszahat, Matyus, on Rubus leaves, 48.29° N, 22.27° E, leg. 21 Jul.

1966 (Kolosvary & Homonnay 1967); Tiszahat, Tiszamogyoros, in tree hollows, 48.32° N, 22.25° E,

leg. 1 Aug. 1966 (Kolosvary & Homonnay 1967); Tiszaorveny, 47.61° N, 20.72° E, leg. Aug. 1968

(Kolosvary 1969); Tiszasiily, 47.39° N, 20.41° E, leg. Aug. 1968 (Kolosvary 1969); Tiszaszederkeny,

47.91° N, 21.08° E, leg. Aug. 1968 (Kolosvary 1969); Tiszatarjan, 47.82° N, 21.01° E, leg. Aug. 1968

(Kolosvary 1969); Tiszavalk, 47.68° N, 20.75° E, leg. Aug. 1968 (Kolosvary 1969); Tiszavarkony,

47.07° N, 20.19° E, leg. Aug. 1968 (Kolosvary 1969); Vencsello, 48.18° N, 21.56° E, leg. Aug. 1968

(Kolosvary 1969); Tisza-volgye, Tiszabercel, Babocsa-szog, 48.16° N, 21.63° E, leg. 20 Jul. 2002

(Lengyel & Muranyi 2006); Tisza-volgye, Tiszabercel, Babocsa-szog, 48.16° N, 21.64° E, leg. 20 Jul.

2002 (Lengyel & Muranyi 2006); Tur River estuary into Tisza River (km 715-744, GPS placed at

Tivadar), 48.06° N, 22.52° E (Avram 1968, type locality of Leiobunum tisciae), 1 <$, 1 §, likely same

locality: Tisza River, left banks, leg. 21 Jul. 1967 (CJM3526, det. by Avram, likely part of the type

series of Leiobunum tisciae , the remainder of specimens not specified, likely in Bucharest Museum but

no data accessible); Zemplen . Erdohorvati, Tolcsva-patak part, 48.29° N, 21.45° E, leg. 3.11.2001

(Lengyel & Muranyi 2006, orig. det. as L. rupestre ); Telkibanya, Cserenko (oberlauf), 48.48° N, 21.35° E,

leg. 30 Sep. 2001 (Lengyel & Muranyi 2006, orig. det. as L. rupestre).

LATVIA: All Latvian records refer to UNILAT, Spungis 2008, if not otherwise indicated. Aizkraukle

Distr., 1 ex., Erberge, rotten birch wood, 56.37° N, 25.02° E, leg. 15 Oct. 2006; 1 ex., Koknese, rotten

deciduous wood, 56.66° N, 25.41° E, leg. 11 Oct. 2003; 2 ex., Krasti, deciduous forest, 56.44° N, 25.92° E,

leg. 15 Oct. 2006; 1 ex., Vesetas paliene fen, on tree trunk, 56.70° N, 25.81° E, leg. 15 Jun. 2005; 4

1 5, Bergi, pie Bukultiem, 56.99° N, 24.28° E, leg. 25 Sep. 1960 (Tumss 1963); Daugavpils Distr .. 1 ex.,

Pilskalnes Siguldina, rotten deciduous wood, 55.98° N, 26.25° E, leg. 15 Oct. 2005; 3 ex., Kraslava

Distr .. Ezernieki, on building, 56.18° N, 27.66° E, leg. 9 Oct. 2005; 6 $ §, Krimulda, 57.19° N, 24.85° E,

leg. 27 Jul. 1937 (Tumss 1963); 2 ex., Kuldlga Distr .. Ivande, on building, 56.99° N, 21.76° E, leg. 22 Oct.

2006; 1 ex., Lunbazi Distr . Mernieki, on wooden building, 57.86° N, 24.51° E, leg. 14 Aug. 2007; 3 ex.,

Madona Distr .. Berzaune, on building, 56.81° N, 26.04° E, leg. 8 Aug. 2007; 2 Moricsala . 57.21° N,

22.14° E, leg. 22 Oct. 1960 (Tumss 1963); Riga Distr .. 3 ex., Salaspils, deciduous forest, 56.87° N,

24.35° E, 21 Sep. 2005; 5 ex., Tumsupe river valley (GPS not accurate), on white alder trunk, 57.01° N,

24.61° E, 22 Aug. 2007; 1 ex., Botanical garden of University of Latvia, greenhouse, pitfall traps,

56.950° N, 24.058° E, leg. 15-29 Nov. 2004; 1 ex., Darzini, on wooden building, 56.87° N, 24.28° E,

leg. 24 Sep. 2006; 4 ex., Nordeki, deciduous park, 56.96° N, 24.17° E, leg. 24 Sep. 2005 (3 ex.), leg.

5 Oct. 2003 (1 ex.); 9 ex., Saldus Distr .. Saldus, Ciecere river valley, on wooden building, 56.67° N,

22.49° E, leg. 22 Oct. 2005; 1 & Sauka, 56.28° N, 25.48° E, leg. 10 Aug. 1937 (Tumss 1963); 1 1 ?,

Sigulda, 57.15° N, 24.85° E, leg. 21 Oct. 1960 (Tumss 1963); 5 $$, Sudas purvs, 57.15° N, 25.00° E,

leg. 1 Oct. 1960 (Tumss 1963); 1 ex.. Talsi Distr .. Dundaga, deciduous park, 57.51° N, 22.36° E, leg. 28 Oct.

2007; 8 ex., Kolka, on wooden building, 57.75° N, 22.58° E, leg. 28 Aug. 2005 (7 ex.), 31 Aug. 2007

(1 ex.); 1 ex., Valmiera Distr .. Kauguri, garden, 57.49° N, 25.44° E, leg. 8 Aug. 2004; Ventspils Distr ..

9 ex., Usma, rotten deciduous wood, 57.22° N, 22.15° E, leg. 13 Aug. 2006.

European Journal of Taxonomy 216 : 1-35 ( 2016 )

NORWAY: Oslo (=Kristiania, coordinates inaccurate), 59.92° N, 10.82° E, type locality of Leiobunum

norvegicum Strand, 1900, 1 $ (Strand 1900); 1 $, 7 juv., Ostlandet, Tistedalen, 59.13° N, 11.44° E, leg.

22 Aug. 1977 (Stol 2010b); 1 ?, Rogaland, Karmoy, 59.28° N, E5.21° E, leg. 1977 (Stol 2010a);

Stavanger, 58.97° N, E5.70° E, leg. ? (SMF, Schonhofer 2005); Vestfold, Botne, 59.482° N, 10.275° E,

leg. 13 Sep. 1964 (MBN-A3575, Stol 2010b).

POLAND: 2 juv., Bialowies (no further data, coordinates inaccurate), 52.67° N, 23.83° E, leg. 15 Aug.

1916 (SMF 1603, Muller 1920); 1 3 , 1 §, Bieszczady Mts., Distr. Mstrzyki Dolne, Mstrzyki Gorne,

“zarosla blehowe” along small stream Terebowiec, 49.10° N, 22.67° E, leg. 25 Sep. 1964 (CJM6368,

I.Z.PAN-Warszawa, 35/64); 11 5?, 14 juv., Bug River Valley, landscape park N Neple, aggregations

under branches close to floor of riverine forest, 100 m, 52.135° N, 23.519° E, leg. 14 Jul. 2010

(AXLS958); Chylonia (formerly Kielau), 54.53° N, 18.44° E (Le Roi 1914); Frombork (formerly Frauen-

burg), 54.35° N, 19.68° E (Le Roi 1914); Janow, 53.47° N, 23.23° E (Wajgel 1874), orig. Z4612, det. as

Leiobunum bicolor ); Masurian Lake District, Lidzbarlc Warminski, 54.12° N, 20.58° E (Stargga 1963);

2 55, 1 juv., Memerki near Wegorzewo, Tilio-Carpinetum, 54.18° N, 21.65° E, leg. 4 Aug. 1983

(SMF36129); Swiekatowo (formerly Westpreuflen, Johannisberg), 53.42° N, 18.09° E (Le Roi 1914);

Warschau vicinity (GPS not accurate), 52.21° N, 21.06° E (Stargga 1963); Wyzynie Lubelskiej, Kazi-

mierz Dolny, 51.317° N, 21.95° E (Stargga 1963).

The following records of Stargga (2004) share the same data: Lysa Gora Mts., different forest com¬

munities (Dentario glandulosae-Fagetum, Tilio-Carpinetum, Pino-Quercetum), xerophilic dry grass¬

land, boulders, caves, buildings, leg. Jul. 1981-Aug. 1985. Data differ in: Kalconin, 50.867° N, 20.929°

E; Miejska Gora, 50.957° N, 20.929° E; NSG Chelmowa Gora, 50.867° N, 21.071° E; NSG Gora

Miedzianka, 50.865° N, 20.361° E; NSG Milechowy, 50.775° N, 20.362° E; NSG Skalki Pieklo pod

Nieklaniem, 51.226° N, 20.642° E; NSG Swiety Krzyz, 50.867° N, 21.071° E; NSG Wykus, 51.047° N,

20.929° E; NSG Zamczysko, 50.777° N, 20.929° E; Szewna, 50.956° N, 21.356° E; 4 33 , 4 5 5 , Lysa

Gora Mts., NSG Lisiny Bodzechowskie, beo Ostrowiec Swietokryskiz, 50.866° N, 21.497° E, leg. 9

Sep. 1981 (SMF, Stargga 2004); 1 3 , 1 5 , Roztoczanski National Park, NSG Nart (coordinates inaccur¬

ate), Dentario glandulosae-Fagetum, 50.60° N, 22.97° E, leg. 15 Oct. 1986 (SMF); 1 5 , Roztoczanski,

R.N., rez. Bulcowa Go’raFB30, leg. 5 Aug. 1981, subadult (SMF).

ROMANIA: 1 5 , Bade Herculane, 44.88° N, 22.42° E, leg. 25 Jun. 1956 (CJM6420); 1 c ?, Bihor Mts.,

valley around Sighistel, 46.52° N, 22.54° E, leg. 26 Sep. 1958 (Avram & Dumitrescu 1969); Bistritz,

Dealul Targului, 47.19° N, 24.50° E (Weiss 1996); 2 33 , CabanaNegoiul, 1540 m, 45.59° N, 24.52° E,

leg. 24 Jul. 1950 (CJM6426); Fogarasch Mts., Balea-Tal, Roter Turm, 45.63° N, 24.26° E (Weiss 1996);

3 ad., Hunedoara, Vulcan, 45.38° N, 23.29° E, leg. 16.11.1999 (CJM6460); 1 3 , 1 ?, Hunedoara, Vul¬

can, 45.38° N, 23.29° E, leg. 16 Nov. 1996 (CJM3770); Klausenburg, Botanical Garden, 46.759° N,

23.586° E (Weiss 1996); 1 3 , 1 ?, Nadrag, 45.65° N, 22.18° E, leg. 30 Aug. 2001 (CJM3768); 1 3 ,

Parang Mountains, Ranca, 45.29° N, 23.69° E, leg. 17 Aug. 1997 (CJM3769); 1 3 , 2 $ $, Per§ani Mts.,

Valea Siclau (coordinates inaccurate, valid for mountains!), 45.90° N, 25.43° E, leg. 4 Aug. 1967 (from

literature); 1 3 , Piatra (Kohat) Mts., Sapan(a (Szaplonca), Brazi valley, 841 m, 47.830° N, 23.743° E,

leg. 2 Sep. 2004 (Muranyi & Lengyel 2006); 8 juv., Piatra (Kohat) Mts., Sapan|a (Szaplonca), mineral

water springs in the lower valley of the Sapan|a (Szaplonca) stream, 408 m, 47.9349° N, 23.6781° E,

leg. 30 Jun. 2005 (Muranyi & Lengyel 2006); 1 juv., Piatra (Kohat) Mts., Sapan(a (Szaplonca), valley of

the Sapan(a (Szaplonca) Stream, 500 m, leg. 30 Jun. 2005 (Muranyi & Lengyel 2006); 1 3 , Prahova,

Sinaia, 45.35° N, 25.55° E, leg. 25 Sep. 1949 (CJM6183, H. Franz, 938); 1 3 , 1 ?, Rodna (Radnai) Mts.,

Sacel (Izaszacsal), Iza spring, 900 m, leg. 22 Sep. 2005 (Muranyi & Lengyel 2006, was described as

L. rupestre but doubtful record); 1 3 , 1 ?, Rodna Mts., Pasul Prislop, 47.609° N, 24.855° E, leg. 25 Jun.

1968 (Avram & Dumitrescu 1969); Saschiz, Keisd, 46.194° N, 24.960° E (Weiss 1996); Sebe§ Olt (GPS

might not be correct), 45.67° N, 24.31° E (Weiss 1996); Suceava, Horodnicu de Sus (Russian translation

MARTENS J. & SCHONHOFER A.L., The Leiobunum rupestre group

and GPS might not be correct), 47.84° N, 25.82° E (Cirdei 1960); Suceava, Magura Mt. 47.32° N,

25.96° E (Ro§ca 1930); Suceava, Rarau Mt., 47.45° N, 25.58° E (Ro§ca 1930); 2 33, Kronstadt, Kapel-

lenberg, 45.635° N, 25.597° E (ZMB12634, orig. det. as L. rupestre ); 2 S3, Sibiu (Hermannstadt),

45.78° N, 24.12° E, leg. 29 Aug. 1896 (ZMB 12636, orig. det. as L. rupestre ).

RUSSIA: Oblast St Petersburg (Leningrad) . Petrodvorec, 59.886° N, 29.896° E (Roewer 1957); Primorje

(formerly GroB Kuhren), 54.94° N, 20.04° E (Ee Roi 1914); Rjabinowka (formerly GroB Raum), 54.83°

N, 20.50° E (Le Roi 1914); Sankt Petersburg (Leningrad), 59.90° N, 30.25° E (Stargga 1978); 1 3, St.

Petersburg, City, (coordinates inaccurate), wall of house, 59.90° N, 30.28° E, leg. 10 Aug. 2003

(CJM3032); Svetlogorsk (formerly Wamicken), 54.90° N, 20.11° E (Le Roi 1914); Swetlogorslc

(formerly Rauschen), 54.89° N, 20.11° E (Le Roi 1914); Alt-Peterhof (St. Petersburg), 6 3 6 $

(SMFRII/1201, Roewer 1957).

SERBIA: 1 3, Bor Dubasnica, Mala Tisnica, 44.200° N, 21.867° E, leg. 12 Aug. 1994 (CJM6363).

SLOVAK REPUBLIC: Cervena skala (Dobsina), 48.82° N, 20.13° E (Silhavy 1981); Dobroc (Brezno

nad Hronom), 48.73° N, 19.69° E (Silhavy 1981, all following records from this reference orig. det as

L. glabrum ); Gornasek (Roznava; used for GPS), 48.66° N, 20.54° E (Silhavy 1981); Kamenica nad Cir

(Kamenica nad Cirochou: GPS), 48.93° N, 22.00° E (Silhavy 1981); Kosice, 48.72° N, 21.25° E (Sil¬

havy (1981) did not discriminate betw. L. rupestre and L. tisciae for this record); Nove Mesto nad

Vahom, 48.75° N, 17.83° E (Silhavy 1981); Pajstun (Bratislava), 48.27° N, 17.08° E (Silhavy did not

discriminate betw. L. rupestre or L. tisciae for this record); Presov, 48.99° N, 21.24° E (Silhavy (1981)

did not discriminate betw. L. rupestre and L. tisciae for this record); Rajecke Teplice, 49.13° N, 18.68° E

(Silhavy 1981); Rohace Mts. (coordinates inaccurate), 49.23° N, 19.72° E (Silhavy 1981); Stubnianske

Teplice (today Turcianske Teplice), 48.86° N, 18.86° E (Silhavy 1981); Sulov, 49.16° N, 18.59° E (Sil¬

havy 1981); Trencianske Teplice, 48.91° N, 18.17° E (Silhavy 1981); Vihorlat Mts. (coordinates inac¬

curate), 48.89° N, 22.13° E (Silhavy 1981); Vratna dolina (Zilina), 49.26° N, 19.03° E (Silhavy 1981).

SWEDEN: Ostergotland (coordinates inaccurate, just for the map!), 58.35° N, 15.51° E (Tullgren 1906a,

b); Vastergotland, Raback, 58.61° N, 13.36° E (Tullgren 1906a, b); Skane County, Lund, 55.70° N,

13.19° E, leg. Aug. 1872 (Tullgren 1906a, b); Smaland (coordinates very inaccurate!), 57.4° N, 15.3° E

(Tullgren 1906a, b).

UKRAINE: Bez. Cernovcv . Ciudei, 48.05° N, 25.62° E, leg. 1935 (Cirdei 1960); Bez. Kiev , Bila Tserkva

(org.: Belaja Cerkov), 49.78° N, 30.13° E (Morin 1934); Bez, Uzgorod . Rachov, 48.05° N, 24.21° E

(Bartos 1939); Bez, Zitomir . Novograd-Volynskij, 50.58° N, 27.66° E (Morin 1934); Chernivets‘ka,

Vijni(a, 48.25° N, 25.19° E (Ro§ca 1930); L‘vovskaya oblast‘ (orig.: Bez. Uvov), Ivano-Frankovo,

49.92° N, 23.73° E (L. Koch 1870); Ternopil, Kremenec, 50.11° N, 25.72° E (Stargga 1978).

Leiobunum rupestre (Herbst, 1790)

AUSTRIA: Vorarlberg : 1 3, Ubersaxen, Baschghohle, 785 m, 47.25° N, 9.67° E, leg. October (Breuss

1995); Salzburg (all records if not otherwise indicated refer to Ausobsky 1987), Bergheim bei Salzburg,

Hochgitzen, 47.850° N, 13.034° E; Buchberg, 47.942° N, 13.092° E; Haunsberg, 47.92° N, 12.99° E;

Heuberg, 47.82° N, 13.11° E; Moosachtal bei St. Georgen an der Salzach, 47.98° N, 12.88° E; Plain,

Plainberg,47.837°N, 13.044°E; Salzburg, Festungsberg, 47.795° N, 13.048°E; Salzburg, Kapuzinerberg,

47.803° N, 13.056° E; Salzburg, Monchsberg, 47.799° N, 13.039° E; Salzburger Becken, Elsbethen,

Elsbethener Hiigel, 47.76° N, 13.10° E, Georgenberg near Kuchl, 47.635° N, 13.161° E; Hellbrunner

Berg, 47.758° N, 13.066° E; Morzger Hiigel, 47.768° N, 13.055° E; Tannberg, 47.973° N, 13.188° E;

Tiefstein gorge near Schleedorf, 47.947° N, 13.158° E; Stubachtal, Weissee, 47.16° N, 12.64° E (Mazzucco

1967); 4 33, 2 55 , Stubachtal, N Enzingerboden, 1420-1460 m, 47.184° N, 12.625° E, leg. 7 Oct.

European Journal of Taxonomy 216: 1-35 (2016)

1966 (CJM496); 11 33, 5 §§, Pinzgau, Kaprun, entrance of Sigmund-Thun-Klamm (Alpl2_04), 800

m, 47.258517° N, 12.738571° E, leg. 10 Sep. 2012 (AXLS1283); Oberosterreich . Weilhartsforst, 48.16°

N, 12.91° E (Ausobsky 1987); 1 3, vicinity ofStifterherberge at Barenstein, 900-950 m, upper coniferous

forest, 48.660° N, 13.991° E (SMF16343/1); 1 3, Steiermarlc Koralpe, Soboth, 900 m, wet slopes near

stream, Salix and Abies, 46.67650° N, 15.08331° E, leg. 30 Aug. 2009 (AXLS277); 1 3, Gesause,

coniferous forest, 47.58° N, 14.64° E, leg. 16 Sep. unknown jear (ZMB12640); Karnten . Baldersdorf,

way to Egelsee, 46.78° N, 13.56° E (Komposch 2008); Saualpe, Lollingbach, 1100 m, 46.55° N, 14.37°

E (Komposch & Gruber 2004); 5 juv., Flaschberg near Oberdrauburg, 46.75° N, 12.94° E, leg. 28 Jul. 1982

(Kofler & Mildner 1986); Klagenfurt, Sattnitz, boulders betw. Bad and Hanswirt, leg. 3 Sep. 1985

(Kofler & Mildner 1986); Klagenfurt, Sattnitz, Hollgraben, 46.49° N, 14.28° E, leg. 10 Sep. 1985 (Kofler

& Mildner 1986); 5 33, 1 ?, near Klebas, road 111, subway, 46.696° N, 12.802° E, leg. 1 Sep. 2012

(AXLS1356); Ruin Obertrixen, 46.702° N, 14.575° E, leg. 19 Aug. 1985 (Kofler & Mildner 1986); St.

Leonhard near Liebenfels, 46.74° N, 14.26° E, leg. 30 Jul. 1985 (Kofler & Mildner 1986); 1 3,

Tropolach, Gailtal, 900 m, 46.62° N, 13.28° E, leg. 30 Sep. 1975 (CJM1411); Ulrichsberg, 46.71° N,

14.31° E, leg. 3 Oct. 1968 (Kofler & Mildner 1986); 6 33, 2 §§, N Graz, Hinterreitenwald, SE Glett-

weber, Glettbach, 870m,47.201°N, 15.428°E,leg. 14 Sep. 2012 (AXES 130 l);EloheTauern, Gossnitztal

(Gossnitz=Goessnitz), 1700 m, 47.031° N, 12.792° E (Komposch & Gruber 1999); Niederosterreich .

Lunz am See, forests arround Untersee, 47.85° N, 15.05° E (Thaler 1963); 1 3, Semmering (M2),

47.63° N, 15.83° E, leg. Oct. 1899 (ZMB 12644); 1 3, Tirol . Innsbruck, in the city, Weierburggasse,

730-800 m, at housewall, 47.278° N, 11.396° E, leg. 27 Sep.-l Oct. 1986 (CJM3523); Innsbruck

surroundings, Picea forests S of Inn River, HuBlhof-Natters-Amras-Igls-Ampass-Hall, 580 m, in moss

(Stipperger 1928); 1 ex., Forellenhof near Lavant, at limestone boulders, 46.79° N, 12.84° E, leg. 7 Sep.

1971 (Kofler 1984); 3 juv., Nikolsdorf, Tal-Trasse, right side ofDrau River, 46.78° N, 12.91° E, leg. 28

Jul. 1982 (Kofler 1984); 1 §, Norsach (Karntner Grenze), 46.78° N, 12.93° E, leg. 28 Aug. 1969 (Kofler

1984); Villgrater Tal (GPS not accurate), 46.80° N, 12.38° E (Werner 1931); 1 $, Virgental, Iselschlucht

near Welzelach, 47.017° N, 12.413° E, leg. 2 Aug. 1978 (Kofler 1984); Windisch-Matrei, 47.00° N,

12.53° E (Dalla-Torre 1882); Barenbadalm above Achensee, 1400-1500 m, caves in Picea forest, 47.42° N,

11.70° E (Stipperger 1928); Hungerburg, 860 m. Sand- and Schotterquarry at border of Picea forest,

47.29° N, 11.40° E (Stipperger 1928); Maldontal, betw. Boden and Bschlabs, 1350 m, Fichten- und

Fohrenwald, 47.30° N, 10.59° E (Stipperger 1928); Stallental, 800-1600 m, Picea forest with Fagus,

47.38° N, 11.69° E (Stipperger 1928); Kaunertal from Kaltenbrunn and Feuchten to Gepatschhaus

(coordinates inaccurate), 1260-1700 m, Picea forest, humid cliffs along stream and path, 47.05° N,

10.75° E (Stipperger 1928); Oberbergtal fromMilders im Stubaito Stocklenalm (coordinates inaccurate),

1100-1600 m, Picea forest, at boulders, 47.12° N, 11.32° E (Stipperger 1928); S St. Anton am Arlberg,

1310 m, cliffs in Picea forest, 47.13° N, 10.27° E (Stipperger 1928); Schwaz, betw. SchloB Freundsberg

and Rodlhiitte, 540-1600 m, Picea forest mixed with Abies and Fagus , under leaf litter, in moss, at

stones and truncs, 47.34° N, 11.71° E (Stipperger 1928); Sellraintal betw. Kematen, Gries and Praxmar,

610-1600 m, humid, shaded cliffs and moss at stream in Picea forest, 47.17° N, 11.14° E (Stipperger

1928); between Pettneu and Malfontal, 1200-1300 m, Picea forest, 47.140° N, 10.343° E (Stipperger

1928); betw. Volders and Volderer Wildbad, 600-1100 m, wet Picea forest with Alnus; 3 33, 47.27° N,

11.56° E (Stipperger 1928); Lienz (GPS not accurate), 46.83° N, 12.76° E, (SMFRII/1184, Roewer

1957); 3 33, 1 ?, Lienzer Dolomiten, Hohenstein (SMFRII/701, Roewer 1957); Burgenland . 1 3,

southern Leithagebirge, „Kurschnergrube” (abandoned chalk quarry) N of Eisenstadt, 435 m, in man

made cavity at steep humid cliff, 47.86° N, 16.52° E, leg. 7 Nov. 1996 (NMW17922, Gruber 2000).

BOSNIA AND HERZEGOVINA: 11 33, 1 ?, Jaice (=Jajce), tufa walls, waterfall, 44.342° N, 17.266° E

(ZMB 12639); 2 33, Kupreska vrata, Bugojno, 1384 m, 44.00° N, 17.31° E (Novak 2005); Travnik,

44.22° N, 17.66° E (Martens 1978); 2 33, Trebevic (National Park and mountain S of Sarajevo,

coordinates accurate), Corylus forest, 43.8° N, 18.45° E, leg. Sep.-Oct. (ZMB 12638); 4 33, 1 juv.,

Vilinska jama cave, 42.73° N, 17.89° E, leg. 1916 (Novak 2005).

MARTENS J. & SCHONHOFER A.L., The Leiobunum rupestre group

CROATIA: 1 Bijeli vrh, Vrhovine, 1074 m, 44.84° N, 15.44° E (Novak 2004b); 1 juv., Kamengrad,

Papuk Mt., beside the Orljava river, 528m, 45.53° N, 17.50° E (Novak 2004b); 5 $$, 2 5?, 2 juv.,

Kamenica Skradnicka, beside the Rudnica brook (WL30), 260 m, 45.20° N, 15.45° E, leg. May 1985

(Novak 2004b); 3 $$, 1 $, Kameniti vrh Mt., Plitvice, 934 m, 44.88° N, 15.57° E (Novak 2004b); 1 <S,

Klanjec, 170 m, 46.05° N, 15.74° E (Babic 1916); 1 Krapina, 290 m, 46.16° N, 15.87° E, leg. Aug.

1914 (Babic 1916); Krizevci, 140 m, 46.03° N, 16.52° E (Daday 1896), orig. det. as Liobunum glabrum );

2 SS, Ledenjaca u Cudinoj uvali cave (WK46), Veliki Javornik, 44.84° N, 15.57° E (Novak 2004b);

Maksimova pecina cave, Jankovac, 657 m, 45.52° N, 17.68° E (Babic 1916); 2 Medvedalc Mt.,

Plitvice, 889 m, 44.89° N, 15.64° E (Novak 2004b); 2 $$, 1 §, E Sveti Jakob, stream valley of river

Medvescak, 600 m, under stones and wet logs near river, 45.884° N, 15.945° E, leg. 9 Sep. 2009

(AXLS345); 1 juv., near Vrilo Rudnice, Kamenica Skradnicka, 260 m, 45.23° N, 15.35° E, leg. Jul. 1987

(Novak 2004b); 1 juv., Ozalj, Karlovac, Kordun, Ozaljska spilja cave, 45.62° N, 15.45° E, leg. 6378

(Novak 2004b); 13 juv., Pecina pri Trzicu (=Pecina kod mlina), Trzic Tounjski, beside the Mreznica

River, 45.23° N, 15.41° E, leg. Jul. 1983 (Novak 2004b); 1 juv., Plitvice (coordinates inaccurate), 677

m, 44.88° N, 15.62° E (Babic 1916); 6 Sagorije, Ogulinsko, Ivancij, hunter house at Smitowo

jezero, 351 m, at large rocks in Fagus forest, 45.188° N, 15.223° E, leg. 7 Sep. 2009 (AXLS319); 1

1 5, Sagorije, Ogulinsko, Ivancij, Sagorska pec, 326 m, at wall in cave entrance, 45.197° N, 15.222° E,

leg. 7 Sep. 2009 (AXLS322); 1 Skrad, 700 m, 45.43° N, 14.91° E, leg. May 1905 (Babic 1916);

1 juv., Sveta gora, 580 m, 45.53° N, 14.67° E (Novak 2004b, Babic det. as O. alpinus ); 1 1

Vilinska jama cave (=Vilina jama, Volina-Volinjska jama, one hour away from Krapina), leg. 11 Aug.

1885 (Jurinac 1886, Novak 2004b), orig. det. as Liobunum religiosum).

_ _ V V

CZECH REPUBLIC: All records originate from Silhavy (1981) so far not otherwise indicated. Sumava,

Boubin, 48.94° N, 13.79° E; Holna (maybe wrong), 49.12° N, 14.87° E; Horni Smrcne, 49.33° N, 15.76°

E; Hruba Skala, 50.54° N, 15.19° E; Jested Mt., 50.73° N, 14.98° E; Jevany, 49.97° N, 14.81° E (Silhavy

did not discriminate betw. L. rupestre or L. tisciae for this record); 2 SS, 1 ?, limestone caves near

Briinn (Brno), leg. 17 Sep. 1971 (CJM982); Klucky (mountain ridge near Novy Bor), 50.78° N, 14.50°

E; Konesin, 49.19° N, 16.04° E; Kunratice, 50.01° N, 14.48° E (Silhavy did not discriminate betw.

L. rupestre or L. tisciae for this record); Kvilda, 49.02° N, 13.58° E; Lamberk (Namesf nad Oslavou),

49.17° N, 16.18° E; Machovo jezero (Dosky), 50.58° N, 14.67° E; Marianske Lazne, 49.96° N, 12.70°

E; Mohelno (Treble), 49.11° N, 16.19° E; Moravsky kras, 49.31° N, 16.70° E; Olsi (Jindrichuv Hradec),

49.16° N, 15.37° E; Pravcicka brana (Hrensko), 50.88° N, 14.28° E; Rokytnice nad Jizerou, 50.73° N,

15.45° E; Rychory, 50.67° N, 15.87° E; Spindleruv Mlyn, 50.73° N, 15.61° E; Tiske skaly (Deem),

50.80° N, 14.04° E; Tremsin (Rozmital), 49.57° N, 13.78° E (Silhavy did not discriminate betw.

L. rupestre or L. tisciae for this record); Vlci kopec, 49.15° N, 16.45° E.

GERMANY: Baden-Wtirttemberg . 2 §?, Nature Reserve Wutachschlucht, Stigelesfels, 48.0166° N,

8.9333° E, leg. 1 Oct. 1980 (CJM1954); 4 SS, Lotenbachklamm, rock face, 47.85° N, 8.3° E, leg. 25

Jul. 1964 (CJM123); 1 Schwabische Alb, Hollochschacht bei Dettingen, 48.513° N, 9.346° E, leg.

2.12.1967 (CJM1198); 2 <$<$, Tiefental, Kreis Niirtingen, Gutenberg, Gubmannshohle, 680 m, 48.543°

N, 9.520° E, leg. 18 Feb. 1962 (SMF13067/2, Dobat 1963); 2 88, 11 Schwarzwald, Triberg,

48.13° N, 8.23° E, leg. Aug. 1959 (SMF11173/12); 5 88, 3 $?, S-Schwarzwald, Schwarzatal S

Schluchsee, Schwarzahalden, 600 m, Bannwald, 47.74° N, 8.19° E, leg. 14 Sep. 2008 (CJM6207); 2

88, 1 juv., Schwarzwald, Titisee, 865 m, 47.89° N, 8.14° E, leg. 29 Sep. 2013 (AXLS1626); Wolfsbrunn

near Heidelberg, 49.412° N, 8.746° E (C. Koch 1871; Le Roi 1914); Bayern . Benediktenwand, 47.65°

N, 11.47° E (L. Koch 1861); Berchtesgaden, Jenner, 800 m, forest, 47.58° N, 13.00° E (Hammelbacher

1986); 1 S, 1 ?, Ramsau / Wachterl, 1000 m, mixed forest, 47.65° N, 12.866° E, leg. 11 Jul. 1986

(CJM2804); 3 SS, Tegernsee, ascend to Wallberg, 700-1000 m, coniferous forest, at cliff, 47.68° N,

11.75° E, leg. 31 Sep. 2006 (CJM5318); near Regensburg (GPS for the city), 48.97° N, 12.13° E (Martens

1978); 4 SS, Bavarian Forest, N Finsternau, Siebensteinkopf, 1130-1200 m, 48.95° N, 13.566° E, leg.

23 Aug. 1992 (CJM4358); 2 Arber, in the cirque-wall (Karwand), 49.10° N, 13.12° E, leg. 19

European Journal of Taxonomy 216: 1-35 (2016)

Aug. 1964 (CJM136); 20 juv., Lusen, Arber, Riesloch Falls, 48.933° N, 13.50° E, leg. 14-17 Aug. 1964

(CJM143); 1 8, 6 juv., Rachelsee, under stones, 48.975° N, 13.402° E, leg. 16 Aug. 1964 (CJM113,

CJM109); 6 88, Riesloch-Falle near Bodenmais, 49.05° N, 13.083° E, leg. 18 Aug. 1964 (CJM101);

1 c?, 1 $, Garmisch-Partenkirchen, Partnachklamm, 850 m, 47.466° N, 11.116° E, leg. 14 Oct. 1978

(CJM1856); 1 ?, NSG Murnauer Moos, 47.64° N, 11.17° E, leg. 1977/78 (SMF33105, Loser et al.

1982); Niirnberg, betw. Bruck and Happburg (coordinates inaccurate), 49.52° N, 11.18° E (L. Koch

1877, as L. bicolor ); Pottenstein, Frankische Schweiz, 49.77° N, 11.40° E, leg. 1 Oct. 1948 (SMF59294,

Schonhofer 2005); 2 88, Waldnaabtal, Oberfranken, 49.6° N, 12.133° E, leg. 22 Aug. 1964 (CJM86).

Hessen . 1 8, Werra-Meibner-Kreis, Meibner, stone cellar near Neuen Schwalbenthaler Erbstollen (No.

4725/58), 580 m, Mittlerer Buntsandstein, 51.2056° N, 9.87 09° E, leg. 25 Oct. 2005 (CJM4802, S.

Zanlcer, WK 211); Thuringen . 2 88, Schwarza Valley near Schwarzburg, 50.633° N, 11.183° E, leg. 28

Sep. 2006 (CJM5392); Sachsen . 1 §, Erzgebirge (Ore Mountains), at Flobgraben betw. Aue and Bockau,

50.55° N, 12.67° E, leg. 21 Nov. 1926 (Buttner 1930); Lausitzer Mts., Neibe Valley betw. Ostritz and

Hirschfelde, 50.97° N, 14.92° E (Hiebsch 1972); 8 88, 1 $, S Konigstein, stream valley before junction

to Cunnersdorf, Biela Stream, 150 m, at stone bridge, 50.903° N, 14.064° E, leg. 9 Sep. 2006 (CJM5293);

7 88, Schona, near border to Czech Republic, 100 m, coniferous and mixed forest, at night at wood and

stones, 50.867° N, 14.217° E, leg. 13 Oct. 2005 (CJM4752); 2 $$, same data (CJM4753).

HUNGARY: Budapest, Mecsek Mts. (coordinates inaccurate), 47.51° N, 19.01° E (Kolosvary 1929,

1964); 10 ex., Geschriebenstein Nature Park, beside road to Velem (St. Veit), NNE Ohaz, WSW Koszeg

(Guns), 400 m, scree in a fresh deciduous forest with maple, linden, hornbeam and beech, 47.22° N,

16.30° E (Komposch 2004); Koszegi-hegyseg, Koszeg, Het-forras, 47.39° N, 16.53° E, leg. 31 Aug.

2005 (Lengyel & Muranyi 2006); Koszegi-hegyseg, Velem, Szerdahelyi-patalc-part, 47.345° N, 16.514° E,

leg. 17 Sep. 2005 (Lengyel & Muranyi 2006).

ITALY: Carnia, 46.37° N, 13.14° E (Caporiacco 1922); 1 8, Friuli-Venezia Giulia . Prov. Udine, Carnic

Alps near Tarvisio, 800 m, 46.50° N, 13.57° E, leg. 1 Oct. 1975 (CJM1456); 2 juv., Moggio Udinese,

Stavoli Cuel Lung alto, Mt. Zovet (UM53), 700 m, Fagetum, pitfall traps, 46.48° N, 13.19° E, leg. 16 Jul.

-20 Aug. 2001 (MFSNU, Novak 2004a); 1 8, SE Arta Therme, path in forest, 46.478° N, 13.069° E, leg.

1 Sep. 2012 (AXLS1349); 1 $, Resia, above Clen, between Casera Coot and Slatina superiore (UM73),

1000 m, Fagetum, pitfall traps, leg. 20 Aug.-26 Sep. 2001 (MFSNU, Novak 2004a); 3 juv., Resia,

between Berdo di Sopra and Rio Malidul (UM73), 1260 m, 46.34° N, 13.29° E, leg. 12 Jun. 2003

(MFSNU, Novak 2004a); Sudtirol . Meran, 46.68° N, 11.17° E (L.Koch 1869, type locality of Leiobunum

glabrum L. Koch, 1869).

POLAND: 3 88, Kitz....? (maybe Kitzelberg; label not legible) Bober Katzbach Mts. (GPS not accurate),

50.96° N, 15.84° E, leg. 6 Sep. 1924 (ZMB12643). Karkonosze Mts. (Stargga 2004).

SERBIA: 1 $, Valjevo, planina Medvenik, 44.24° N, 19.63° E, leg. 1 Aug. 1998 (CIK1337); 1 8,

Valjevo, planina Medvenik, Drenaicka pecina, leg. 20 Oct. 1999 (CIK1336).

SLOVENIA: Alto Isonzo, Monte Merzli, 46.22° N, 13.70° E, leg. Nov. 1915 (Marcellino 1973); 1 8,

Celje (Cilli), deciduous forest, 46.24° N, 15.26° E, leg. 25 Oct. (ZMB12635); 7 88, HudaLuknaja Cave,

TisnickMt. SE Slowenj Gradec, 46.48° N, 15.07° E, leg. 16 Oct. 2004 (CJM4203); 3 $ HudaLuknaja

Cave, TisnickMt., SE Slowenj Gradec (GPS S of city), 46.48° N, 15.07° E, leg. 16 Oct. 2004 (CJM4204);

15 88, 4 $ $, Julian Alps, road to Mount Mangert, upper end of village Strmec na Predelu, 1013 m,

46.41769° N, 13.60833° E, leg. 28 Aug. 2009 (AXLS264); 4 88, Karawanken, road to Loibl Pass,

650 m, 46.40° N, 14.28° E, leg. 2 Oct. 1975 (CJM1445); 6 88, 1 ?, Ljubljana, small forest at northern

city limit, 46.10530556° N, 14.51822° E, leg. 3-4 Sep. 2012 (AXLS1389); 9 88, 1 ?, Pohorje Mts.,

Mislinja, locality Glazuta, 1013 m, stream valley, on cliff-faces in coniferous forest, 46.480° N,

15.280° E, leg. 26 Aug. 2009 (AXLS237); 3 88, Savinje valley, Erjavceva Jama, entrance zone, 700 m,