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Redescription of Trogulus nepaeformis (Scopoli, 1763), an often misinterpreted harvestman species from the south-eastern fringe of the Alps (Opiliones: Trogulidae)

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  • Faculty of Natural Sciences and Mathematics, University of Maribor

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In terms of taxonomy and systematics the harvestman genus Trogulus Latreille, 1802 is one of the most complex arachnid taxa in Europe. Although many problems have been resolved and a number of new species were described since 2008, so far, the diversity within several species subgroups within the genus has not been satisfactorily disentangled. One such understudied subgroup is the nepaeformis species-group of medium-sized troguli, partly because we are still missing a thorough reconsideration of the first described species. Trogulus nepaeformis Scopoli, 1763 is at the same time the first harvestman species described from the territory of Slovenia. Here we provide the redescription of Scopoli’s species to set a firm base for revising the whole nepaeformis species-group.
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81
Redescription of
Trogulus nepaeformis (Scopoli, 1763),
an often misinterpreted harvestman species
from the south-eastern fringe of the Alps
(Opiliones: Trogulidae)
Ponovni opis
Trogulus nepaeformis (Scopoli, 1763),
pogosto napačno vrednotene vrste
suhih južin z obronkov južnih Alp
(Opiliones: Trogulidae)
Ton e NOVA K1, Axel L. SCHÖNHOFER2, Ljuba SLANA NOVAK3,
Peter KOZEL1,4, Saška LIPOVŠEK1,5,6, Jochen MARTENS7,8
Abstract
In terms of taxonomy and systematics, the harvestman genus Trogulus Latreille, 1802 is one
of the most complex arachnid taxa in Europe. Although many problems have been resolved and
a number of new species described in last decennia so far, the diversity within several species
subgroups within the genus has not been satisfactorily disentangled. One such understudied
subgroup is the nepaeformis species-group of medium-sized troguli, partly because we are still
Trogulus nepaeformis Scopoli,
           

1 Department of Biology, University of Maribor, Koroška 160, SI-2000 Maribor, Slovenia, tone.novak@guest.
um.si ; peter.kozel@um.si
2 Naturhistorisches Museum und Landessammlung für Naturkunde Rheinland-Pfalz, Reichklarastraße 10,
D-55116 Mainz, Germany dr.axel.schoenhofer@stadt.mainz.de
3 Ozare 31, 2380 Slovenj Gradec,
4 Karst Research Institute, Research Centre of the Slovenian Academy of Sciences and Arts, Titov trg 2,
6230 Postojna, Slovenia
5 Faculty of Medicine, University of Maribor, Taborska ulica 8, SI–2000 Maribor, Slovenia, saska.lipovsek@um.si
6 Faculty of Chemistry and Chemical Engineering, University of Maribor, Smetanova ulica 17, SI–2000 Maribor,
Slovenia
7 Johannes Gutenberg-Universität, Institut für Organismische und Molekulare Evolutionsbiologie (iomE),
D-55099 Mainz, Germany, martens@uni-mainz.de
8 Senckenberg Research Institute, Arachnology, D-60325 Frankfurt am Main, Germany
Tone NOVAK, Axel L. SCHÖNHOFER, Ljuba SLANA NOVAK, Peter KOZEL, Saška LIPOVŠEK, Jochen MARTENS:
Redescription of Trogulus nepaeformis (Scopoli, 1763) ... /Ponovni opis Trogulus nepaeformis (Scopoli, 1763) ...
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SCOPOLIA No 105 – 2023
the whole nepaeformis species-group. Trogulus nepaeformis is characterized by short, stout
and evenly dorsally bent stylus of glans penis with blunt tip, and acinous receptaculum seminis
consisted of eight balloon-shaped and one short-tubular vesicles.
Key words: Arachnida, Idrija, nepaeformis species-group, Slovenia
Izvleček
Trogulus Latreille, 1802,
med najbolj zapletenimi taksoni pajkovcev v Evropi. V zadnjih desetletjih so bile razrešene

             
nepaeformis, deloma zato, ker prva opisana vrsta iz skupine, Trogulus nepaeformis Scopoli,


vrste z namenom, da pripravimo verodostojno osnovo za revizijo celotne skupine nepaeformis.
Za T. nepaeformis
osmih balonastih in ene kratkocevaste ampule.
Ključne besede: Idrija, pajkovci, skupina vrst nepaeformis, Slovenija
83
Introduction
Trogulidae Sundevall, 1833 is a family within the harvestman suborder Dyspnoi Hansen &
Sørensen, 1904, and involves the genera Anarthrotarsus Šilhavý, 1967, Anelasmocephalus Simon,
1879, Calathocratus Simon, 1879, Koniotis Roewer, 1940 and Trogulus Latreille, 1802 (
1978;  2009, 2013). All trogulids are litter- and soil-dwelling and display a conserved
morphology adapted to the habitat and, consequently, are prone to a large degree of cryptic diversity
( 2009, 2013). Additionally, and in Trogulus in particular, several external shapes
and structures are hard to research. This is because all trogulids produce a presumably tanning

particles onto the surface, which hardens into a soil-encrusted, insoluble mail ( et al. 2016).

( &  2009), a time-consuming procedure, which sometimes causes damage

            Trogulus lineage
B sensu  &  (2010), which the nepaeformis group is part of. This lineage
comprises nearly all Trogulus taxa inhabiting central, southern and south-eastern Europe and the
Near East, in contrast to species belonging to the lineage A from the western Mediterranean area
( &  2010). To complicate matters, two or three medium-sized troguli species
co-occur in many localities (e.g.,  1983, 1984). Thus, although the taxonomic position of
Trogulus in Central Europe seems straightforward, it is actually extremely complex ( 1988).
In Slovenia, beside only one Anelasmocephalus species, A. hadzii Martens, 1978, a dozen
of Trogulus species occur, about half of which are medium-sized, mostly belonging to the T.
nepaeformis and T. squamatus species-groups sensu  &  (2010) . Ta xon om ic al
problems within these groups have partly been dealt with morphologically by  (1802),
 (1839),  (1903),  (1971),  (1978),  (1978),  (1980),
 (1983, 1984),  &  (1988) and  et al. (1998). Beside T. nepaeformis
(Scopoli, 1763), the following presently valid species of the nepaeformis and squamatus groups
have been recognized in Slovenia: Trogulus tingiformis C. L. Koch, 1848, T. closanicus Av ram,
1971, T. martensi Chemini, 1983 and T. cisalpinus Chemini & Martens, 1988.
Combining mor phological and genetic information provided a progress i n Trogulus taxonomy
in the last two decennia, and revealed a much-underestimated cryptic diversity within the
genus ( 2008;  &  2010). This approach allowed resolving
a number of open taxonomic and distributional problems ( 2008;  &
 2008, 2009, 2010;  et al. 2013). In Slovenia, syntopic occurrence of up to
at least three Trogulus forms of comparable sizes is quite usual.
For many described species,  (1763) produced perfect drawings allowing unambiguous
recognition. Unfortunately, he did not provide either drawings or the information on locus typicus
of his Acarus nepeformis, i.e., Trogulus nepaeformis
burned in his private house ( 1972;  1977;  et al. 1999), and there has
been no indication of type material being deposited elsewhere and must be considered lost since.
Thus, the only available information on Acarus nepeformis remains his textual description. At
the time being, the name T. nepaeformis is in ample use in central Europe to denote the species
allegedly described by Scopoli and redescribed under this name by  (1978) and 
(1983, 1984). Apparently, misinterpretation of Scopoli’s intention often occurred (e.g.,  1971,
 1978,  2008), resulting in a very broad species concept of “T. nepaeformis”,
including other, partly undescribed species. For a credible approach to the remaining cryptic
diversity within the T. nepaeformis complex it is critical to resolve the identity of Scopoli’s species,
Tone NOVAK, Axel L. SCHÖNHOFER, Ljuba SLANA NOVAK, Peter KOZEL, Saška LIPOVŠEK, Jochen MARTENS:
Redescription of Trogulus nepaeformis (Scopoli, 1763) ... /Ponovni opis Trogulus nepaeformis (Scopoli, 1763) ...
84
SCOPOLIA No 105 – 2023

crucial to resolve the central European Trogulus taxonomy ( &  1988).
In this contribution we aim 1) to establish the terra typica and locus typicus of T. nepaeformis,
          
similar, medium-sized troguli, and 3) to redescribe this species to provide a clear view on this

Materials and Methods
             
original description of Acarus nepeformis Scopoli, 1763 ( 1763) to provide any cues for
establishing the terra typica, or, even more precise, the locus typicus. Besides, we reconsidered
studies of the authors who evaluated Scopoli’s life and work ( 1972; 
1977;  1978;  1999;   et al. 1999) with special attention to any further
indications, which might help in identifying the type locality. Given that Scopoli collected A.
nepeformis for description in a beech forest, we also considered the distribution of the European
beech (Fagus sylvatica L.) at potential localities in the area ( 1986;  et al.
1987;  1998;  et al. 1999;  &  2000). A further
relevant cue is the measure used by Scopoli in measurements of his specimens ( 1763;
 1985;  1999). Finally, we took into account the abundance and general
appearance of syntopic Trogulus species at the potential type locality.
We analysed the harvestman fauna within the radius of 10 km around Idrija, Scopoli’s town
of residence, to provide information on the candidate species and selected the most credible
species. For the redescription, external morphology and genitalia were studied using a Leica
M205 C mounted with a digital camera Leica MC190 HD (Leica Camera AG, Germany),
with Leica Application Suite X 5.1 software. Besides, we used Nikon Eclipse E800 compound
microscope (Nikon, Japan), mounted with a digital Net DN100 camera and processed with
NIS Elements ver. 4.20 software. Transmission and illumination from above were combined

focal planes were stacked using Helicon Focus ver. 6.2.2. Images were manipulated for optimal
quality using various applications in GIMP ver. 2.10.8. Drawings were made under an Olympus
CH30 microscope (Olympus, Japan), using a drawing tube. For observation, the specimens
were preserved in glycerol. Measurements are in millimetres, if not otherwise indicated.
In the redescription, we use the following abbreviations ( 1978,  2008,
 &  2009):
BS basitarsus (basal, i.e., proximal tarsal article or segment), Ch chelicera, Cx coxa, DS
distitarsus (distal tarsal article), Fe femur, Mt metatarsus, Pa pedipalp, Pe penis, Pt patella, Rec
sem receptaculum seminis, Ta tarsus, Ti tibia, Tr trochanter.
Body length: total length from front cap to rear end of opisthosoma
Body width: maximum width of opisthosoma
Coxa II medial interdistance: distance between promedial sides of left and right coxa II
Coxa II lateral interdistance: distance between prolateral sides of left and right coxa II
Coxa IV medial interdistance: distance between retromedial sides of left and right coxa IV
Coxa IV lateral interdistance: distance between retrolateral sides of left and right coxa IV
Eye interdistance: distance between outer borders of left and right lens
Hood (head cap) length: distance between hood tip and anterior border of eyes
Hood width: maximal width of hood
85
Results
Identication of the locus typicus
 (1763) wrote his volume “Entomologia carniolica” while residing in Idrija (1754–
1769), western Slovenia (in older literature Idria in the Duchy of Carniola) (K 2023).
Despite his collecting journeys to many places in Carniola ( 1972), he putatively
described most species from Idrija and its close vicinity ( 1972;  1977;
 1978;  1999;   et al. 1999). Nowadays, troguli are relatively abundant in
the area and it was most probably the case in the past centuries; we thus consider Idrija and the
close vicinity as terra typica of Acarus nepeformis.
Identication of most credible candidate species
              
harvestman species have been recorded (own unpublished data):
CYPHOPHTHALMI
Sironidae
Cyphophthalmus duricorius Jos eph, 1868
(PALPATO R ES)
EUPNOI
Phalagiidae
Amilenus aurantiacus (Simon, 1881)
Dasylobus graniferus (Canestrini, 1871)
Gyas annulatus (Olivier, 1791)
Lacinius dentiger (C. L. Koch, 1847)
Lacinius ephippiatus (C. L. Koch, 1835)
Lophopilio palpinalis (Herbst, 1799)
Mitopus morio (Fabricius, 1799)
Opilio dinaricus Šil havý, 1938
Opilio parietinus (De Geer, 1778)
Phalangium opilio Li nnae us, 1758
Rilaena triangularis (Herbst, 1799)
Sclerosomatidae
Astrobunus helleri (Ausserer, 1867)
Astrobunus laevipes (Canestrini, 1872)
Leiobunum limbatum L. Koch, 1861
Leiobunum rupestre (Herbst, 1799)
Nelima sempronii Szalay, 1951
DYSPNOI
Dicranolasmatidae
Dicranolasma scabrum (Herbst, 1799)
Nemastomatidae
Carinostoma carinatum (Roewer, 1914)
Histricostoma dentipalpe (Ausserer, 1867)
Tone NOVAK, Axel L. SCHÖNHOFER, Ljuba SLANA NOVAK, Peter KOZEL, Saška LIPOVŠEK, Jochen MARTENS:
Redescription of Trogulus nepaeformis (Scopoli, 1763) ... /Ponovni opis Trogulus nepaeformis (Scopoli, 1763) ...
86
SCOPOLIA No 105 – 2023
Mitostoma chrysomelas (Herm ann, 1804)
Nemastoma bidentatum schmidti Novak, Raspotnig et Slana Novak, 2021
Paranemastoma quadripunctatum (Perty, 1833)
Trogulidae
Anelasmocephalus hadzii Martens, 1978
Trogulus closanicus Avram, 1971
Trogulus martensi Chemini, 1983
Trogulus nepaeformis (Sc opoli, 1763)
Trogulus oltenicus Avram, 1971
Trogulus nepaeformis sensu Chemini 1983, 1984
Trogulus sp. gr. tricarinatus Linna eus, 1767
Trogulus tingiformis C. L. Koch, 1848
Among troguli, two small-sized species: T. oltenicus and T. sp. gr. tricarinatus belong to
the tricarinatus species-group; one medium-sized: T. closanicus belongs to the squamatus
species-group, and one large: T. tingiformis and three medium-sized troguli: T. martensi, T.
nepaeformis s. s. and T. nepaeformis sensu  (1983, 1984) belong to the nepaeformis
species-group (cf.  2008;  &  2010). Among these, T.
closanicus preferred humid, deep organic habitats, T. tingiformis humid, deep humus and
stony habitat, and T. martensi and T. nepaeformis sensu Chemini inhabit open woodlands and
scrublands and are absent in beech forests. According to the ratios of the potential candidate
species in syntopy (nepaeformis : closanicus : tingiformis : nepaeformis   
100 : 4 : 2 : 2), we selected the most abundant species in the beech forests at Idrija as the most
credible candidate to identify T. nepaeformis as intended by Scopoli.
Nepeformis. - long.
lin. 4. Diagn. Color & Facies Nepae Cimicoidis. Totus punctis eminentibus confertis scaber.
Reperti non semel inter Muscos, ad Fagorum radices. Oblongus, depressus instar folii, coloris
terrei, iners valde. Antennae moniliformes. Pedes antici & medii breviores. Os ovatum. Pedes
secundi & postici corpore longiores.” (In translation: “1070. ACARUS Nepeformis. – Length
4 lines. Diagnosis. Colour and appearance as in Nepa Cimicoides. Entirely coarse because
of tubercles. Found several times in moss at beech roots. Oblong, leaf-like depressed, soil-
coloured, quite motionless. Antennae necklet-shaped. Fore and middle legs shortest. Mouth
oval. Second and hind legs longest.”). This description is by far incomplete in terms of today’s
nomenclatural code. Hypothetically, it might refer to either any medium-sized Trogulus species
or to small individuals of T. tingiformis, what deserves a detailed analysis.
In his descriptions,  (1763; Explicatio) used the Paris line for size indication, showing
lines in three Paris twelfths – linea; t res unciae Parisinae ( 1999). This is somehow astonishing
because the use of either the Vienna line or the Southern line would be expected. This is for the
reason that the Vienna line was derived from the “Vienna foot, ‘Wiener Fuss’, which was in use
in Central Europe ongoing from about 1760, while the Southern line was in use at the northern
Adriatic coast, the nowadays western and central Slovenian inland, Istria and Friuli (
1985: Küstenland, Krain, Istrien, Friaul). The printed Par is line in  (1763) measured 2.24 mm
( 1999), while the Vienna line measured 2.195155 mm, and the Southern line 2.063305 mm
( 1985). Since Scopoli measured specimens very precisely, often citing quarters,
thirds and halves of the line (e.g., 5 2/3 for Nepa cinerea), his measurements were accurate at about 0.5
mm, and his Acarus nepeformis measured about 9.0±0.5 mm. These measures comply with females
of only one, the most abundant Trogulus species found in Idrija and its vicinity.
87
Accordingly, the following assumption is required. Neglecting the sexual dimorphism and not
taking genitalia into account in his description, Scopoli cited measures of larger individuals –
as usually applied by ancient entomologists and arachnologists – females in this case. Scopoli’s
comparison of the colour and the habitus of T. nepaeformis with the Saucer bug, Ilyocoris
cimicoides (Linnaeus, 1758), Naucoridae ( et al. 1994), in  (1763) quoted sub
Nepa, refers to the anterior portion of this water bug and the hood of troguli; although the shape

the name antennae for the hood horns, and likely interpreted the space below the hood with
chelicerae and pedipalps as the “mouth".
              
 (1977),  (1978),  (1999) and   
that T. nepaeformis had been found several times in moss between beech roots is a case of
 1999). Nevertheless, it is obvious that Scopoli inspected

located close to a beech forest. Nowadays, the beech forests are restricted to the northern and
some other very steep slopes at Idrija. The nearest Fagetum
house, as several buildings have been built in between in the last decennia.
For wood needs, the mercury mine in Idrija owed and exploited a large forest area in Idrija
and its surroundings in the past, which included also clear-cuts in the 18th and 19th centuries,
while moderate steep slopes were already used as meadows and pastures in the 18th century
               
Nowadays, the beech is the dominant species in various habitats from the lowland to the
highest localities in nearly all climatogenous forest phytocenoses in the region ( 1986;
 1998;  &  2000). In this respect, the area is in
harmony with Scopoli’s microhabitat description.
            
exceeding the abundances of the other similar species around Idrija, and reasoning that Idrija is
most likely the type locality, we conclude that T. nepaeformis s. s. described by Scopoli is the
species redescribed below.
Redescription of Trogulus nepaeformis (Scopoli, 1763)
Class Arachnida Lamarck, 1801
Order Opiliones Sundevall, 1833
Suborder Dyspnoi Hansen & Sørensen, 1904
Family Trogulidae Sundevall, 1833
Genus Trogulus Latreille, 1802
SCOPOLIJEV PLOŠČEK, Trogulus nepaeformis (Scopoli, 1763)
Etymology
Scopoli denoted the appearance of the species similar to the water bug Ilyocoris cimicoides
(Linnaeus, 1758) ( 1763, sub Nepa), while the Slovenian name is selected to denote the
Trogulus species described by Scopoli.
Diagnosis
Medium-sized Trogulus species of the nepaeformis species-group sensu 
(2008) and  &  (2010) with ~ roundly shaped head cap (hood), small eyes
Tone NOVAK, Axel L. SCHÖNHOFER, Ljuba SLANA NOVAK, Peter KOZEL, Saška LIPOVŠEK, Jochen MARTENS:
Redescription of Trogulus nepaeformis (Scopoli, 1763) ... /Ponovni opis Trogulus nepaeformis (Scopoli, 1763) ...
88
SCOPOLIA No 105 – 2023
bordered with black ring; with short, stout, evenly dorsally bent stylus of glans penis with blunt
tip; receptaculum seminis acinous, consisted of eight balloon-shaped and one short tubular

Remark: Until the detailed revision of related medium-sized troguli is provided, this diag nosis
should be considered contemporary.
Material examined
Neotype

2000, litter sift collection, L. Slana Novak and T. Novak leg.; PMSL-Opiliones-TN 234/2000.
(PMSL = Prirodoslovni muzej Slovenije, Ljubljana = Slovenian Museum of Natural History,
Ljubljana; this material is a part of the Central Collection of Opiliones at PMSL; TN: Tone

Further material of the species considered

        

 






Novak and T. Novak leg. (Coll. J. Martens CJM 3832 in SMF = Senckenberg Forschungsinstitut

Further specimens will be deposited at the museums of natural history in Vienna, Austria
and Geneva, Switzerland.
Redescription
Male, neotype
Body 7.24 long, 2.85 wide, oblong-oval (dorsal view), strongly dorso-ventrally depressed
(~ half as high as wide), with evenly round-arched head cap (hood) in front of prosoma–hood
lateral indention, 0.65 long, 1.17 wide; distance between hood apex and hood–prosoma lateral
indention 0.85; hood consisted of two latero-anterior prosoma projections (horns, branches)
with long, bristle-tipped, papillate tubercles on medial borders, protruding radially and densely
covering space between horns, and shorter papillate tubercles on lateral horn borders protruding
ventrally, declining their size distally; eye mound relatively low, ~ 2.00 times as wide as long,
0.31 times as high as long, eyes on mound halves separated by intermediate depression; eyes
small, lens diameter 0.11, bordered with blackish ring, eye interdistance 0.65, interocular

giving coarse appearance, and sparse short bristles; heavily earth-incrusted.
Chelicerae proportionally large, Ch basal article ~ 0.2 times as long as body, proximal third
of basal article stout, with large lateral and smaller medial humps inferiorly, wrinkled medially,
89
distal portion abruptly narrowing to ¾ height, smooth; distal article widening distally until




Pedipalps very short, leg-shaped, slender, Pa-Fe proximal half widening distally, distal half
with parallel dorsal and ventral margins (lateral view). Article lengths in Table 1.
Penis 2.23 long, strongly dorso-ventrally depressed, base widest, proximally roundly
indented; truncus, except distal ~ sixth, hard-chitinous, truncus proximal half slightly tapering
distally, followed by portion with ~ parallel margins, and distal, thin-chitinous portion slightly
tapering up to glans; thin-chitinous portion with velum-shaped, shallow fold dorsally, extending
obliquely anteriorly around truncus until ending ventrally para-medially (dorsal view); glans
hard-chitinous, conical, ~ 0.10 times as long as truncus, stylus stout, at base ¾ as wide as glans
terminally, ~ 0.6 times as long as glans, both ventral and dorsal stylus margins ~ arch-bent
dorsally, glans tip blunt.
Legs relatively short, robust, subcylindrical, blackish in old individuals, greyish-brown in

more widely spaced than on body, with relatively dense, long bristles, longest ~ 0.12. Article
lengths in Table 1. Tarsal formula (as in all Trogulus species in both sexes): I 2, II 2, III 3, IV 3. Mt

Cx IV retro-medial interdistance: 2.86, Cx IV retro-lateral interdistance: 3.68.
Fema le (n=1)

      
length 0.57.

Ovipositor 1.72 long, Rec sem acinous, consisted of eight balloon-shaped and one short-
tubular vesicles.
Legs.
Table 1. Trogulus nepaeformis (Scopoli, 1763), neotype, male (female in parentheses). Length of appendage
segments in millimetres in individuals from Idrija, Slovenia.
Trochanter Femur Patella Tibia Metatarsus Tarsus Tot al len gth
Pedipalp 0.33 (0.36) 0.67 (0.73) 0.32 (0.32) 0.50 (0.56) 0.37 (0.38) 2.19 (2.35)
Leg I 0.74 (0.79) 1.67 (1.81) 0.94 (0.98) 1.20 (1.21) 1.11 (1.26) 0.64 (0.62) 6.30 (6.67)
Leg II 0.67 (0.92) 2.92 (3.19) 1.14 (1.75) 1.67 (1.32) 2.03 (2.24) 2.02 (1.75) 10. 45 (11.17 )
Leg III 0.72 (0.77) 1.93 (1.93) 0.99 (1.10) 1.37 (1.53) 1. 52 (1.67) 0.77 (0.73) 7.30 (7.73)
Leg IV 1.00 (1.12) 2.40 (2.88) 1.24 (1.27) 1.90 (1.98) 2.09 (2.23) 0.86 (0.82) 9.49 (10.30)
Tone NOVAK, Axel L. SCHÖNHOFER, Ljuba SLANA NOVAK, Peter KOZEL, Saška LIPOVŠEK, Jochen MARTENS:
Redescription of Trogulus nepaeformis (Scopoli, 1763) ... /Ponovni opis Trogulus nepaeformis (Scopoli, 1763) ...
90
SCOPOLIA No 105 – 2023
Plate I: Trogulus nepaeformis (Scopoli, 1763), neotype, male. 1 Body, dorsal view. 2, 3 Hood. 4. Body, lateral
view. 5 Scutum microsculptures. 6–8 Chelicera: medial, lateral, medial views. 9–11 Pedipalp: medial, lateral,
medial views. 12 Leg II: metatarsus and tarsus. 13. Leg II, femur armament. 14, 15 Penis, dorsal, lateral views.
16, 17 Glans lateral view. 18, 19 Glans, dorsal view. Photo T. Novak & P. Kozel, drawings T. Novak.
91
Plate II: Trogulus nepaeformis (Scopoli, 1763), female. 1–3 Body; dorsal, ventral, lateral views. 4 Hood. 5
Scutum microsculptures. 6, 7 Chelicera: medial, lateral views. 8, 9 Pedipalp: medial, lateral views. 10, 11
Ovipositor. 12, 13 Receptacula seminis.
Tone NOVAK, Axel L. SCHÖNHOFER, Ljuba SLANA NOVAK, Peter KOZEL, Saška LIPOVŠEK, Jochen MARTENS:
Redescription of Trogulus nepaeformis (Scopoli, 1763) ... /Ponovni opis Trogulus nepaeformis (Scopoli, 1763) ...
92
SCOPOLIA No 105 – 2023
Remarks on taxonomy
Acarus Nepeformis Scopoli, 1763 was placed in the correct arachnid order, Opiliones, by
 (1802) who in accordance with the present edition of the Code Art. 24.2.3 (ICZN
1999) acted as the First Reviser and renamed it as Trogulus nepaeformis (Scopoli, 1763)
( 2013). Trogulus nepaeformis s. s. is very similar to an until now unresolved
number of additional species belonging to the nepaeformis species-group sensu 
(2008) and  &  (2010), which have constantly been cited under this name.
The revision of the group is cu rrently at the very beginning. So far, only few authors authentically
refer to T. nepaeformis s. s. in the territory of Slovenia ( 1763;  1997;  &
 2000;  2003;  &  2004;  et al. 2006, 2007; 
et al. 2013a, b;  et al. 2021). Although body length and width are considerably
variable, on the one hand, the genital morphology (penis, receptacula seminis) and measures

the species. However, any synonymy within the nepaeformis species-group cannot be provided
until the revision of the group is completed. For example, a number of populations from other
areas are not congruent with this redescription presented here, especially in various parts of the
Alps, also varying with elevation. Furthermore, populations on the Balkan Peninsula are to be
considered, where Trogulus taxonomy in other groups was shown to be exceedingly complex
( &  2008;  et al. 2013).
Table 2. Trogulus nepaeformis (Scopoli, 1763) variability in body length and width, lengths of Mt II, BS, DS and
whole Ta II in millimetres, and Mt II/Ta II ratios in individuals from Idrija, Slovenia. Mean±StDev (min–max).
Body length Body width Mt II BS DS Ta II Mt II/Ta II

(n=9) 7.3 0.24
(6.94 7.67 ) 2.90±0.09
(2.77–3.07) 2.02±0.05
(1.93–2.01) 0.82±0.03
(0.78– 0.88) 1.23±0.03
(1.18–1.26) 2.05±0.04
(1.98 –2 .11) 0.99±0.03
(0.9 4–1.03)

(n=10) 8.38 ±0.21
(8.05–8.67) 3.30±0.26
(2.82–3.63) 2 .18± 0.08
(2.11–2.32) 0.77±0.03
(0.74–0.85) 1.11±0.05
(1.03 –1.22) 1.89±0.04
(1.85–2.00) 1.15±0.05
(1.10–1.23)
Distribution
Though our comparative material from the environments of Idrija is quite rich and amounts

to date. Apparently, T. nepaeformis covers a rather small area at the fringes of the south-eastern
Alps. For the time being, its precise distributional area cannot be established prior to a revision
of all medium-sized troguli within the eastern Alps and the north-western Balkans.
Ecology
Like other ground-dwelling species, T. nepaeformis live in litter, soil, stone accumulations
and similar spacious microhabitats, at daytime hiding under stones, dead wood on the ground, in

and dolomite, as it is most abundant there. In nature, individuals have been found to feed on
enchytraeids, small lumbricids and soft-chitinous, mostly dipteran larvae (T. Novak & L. Slana
Novak, unpublished observations), but they most probably feed also on snails, as do troguli of a
related species of the nepaeformis species-group ( 1953, sub T. nepaeformis).
93
Discussion
Sharing the Alpine, central European, Dinaric, Sub-Mediterranean and Sub-Pannonian
             
biogeographical regions. There are more than ten Trogulus species in the country ( 1931;
 1984;  2000;  &  2000;  et al. 2006; 
2009;  &  2009;  &  2011; own unpublished data).
However, we still do not have a clear idea about the number of valid species in Slovenia, not to
speak about valid names.
The epithet nepaeformis is used as a collective name for a multitude of European medium-
sized troguli with the ratio of ‘distal segment: basal segment’ of tarsus II approximately 1.5: 1
( 2009). It is important that species of the nepaeformis species-group are similar in

( 2009). Moreover, it is encouraging that comparable genetic distances coincide
with morphospecies ( 2009). The assumption that Scopoli collected specimens
of T. nepaeformis in Idrija – his living and working place at that time – was essential to focus
preparation of our redescription. Selecting by far the commonest species of the four medium-
sized troguli around Idrija as credible candidate of Scopoli’s species was consequently much

most qualifying conditions of article 75.3. of the ICZN. Still, delineation of this and related
species is an ongoing process (article 75.3.2), and the redescription of T. nepaeformis serves
as a base for revising medium-sized troguli and for considering the synonymy within the
nepaeformis species-group.
Acknowledgements
Konrad Thaler (†) helped with literature support and discussion on the measures used by
Scopoli and C. L. Koch. Andrej Gogala helped with the discussion on Ilyocoris cimicoides.
              
Idrija in past centuries. Tone Wraber (†) critically read an earlier version of the manuscript
and provided insightful suggestions and corrections. Christian Komposch provided a peer
reviewing with many helpful suggestions and corrections. We are indebted to all of them. For
a few years, T. Novak was partly supported by the Slovene Ministry of Higher Education,
Science and Technology within the research programme Biodiversity (grant P1-0078). P. Kozel
acknowledges the Karst Research programme (research core funding No. P6–0119). This
study was partly supported by the Project “Development of research infrastructure for the
International competitiveness of the Slovenian RRI space – RI-SI-LifeWatch” (the operation

European Union from the European Regional Development Fund). Over the years, J. Martens
was sponsored by Feldbausch-Stiftung and Wagner-Stiftung at Fachbereich Biologie of Mainz

Tone NOVAK, Axel L. SCHÖNHOFER, Ljuba SLANA NOVAK, Peter KOZEL, Saška LIPOVŠEK, Jochen MARTENS:
Redescription of Trogulus nepaeformis (Scopoli, 1763) ... /Ponovni opis Trogulus nepaeformis (Scopoli, 1763) ...
94
SCOPOLIA No 105 – 2023
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Article
Full-text available
Trogulus falcipenis, spec. nov., the smallest hitherto known species of the genus, is described. This trogulid is characterized by its small body size (< 4,7/ < 5,3 mm), large interocular distance, dark-brown chelicerae and in males, by the short penis (< 1,25 mm), the rod-shaped truncus and the sickle-shaped stylus. Further morphological differences compared to T. tricarinatus (Linnaeus, 1767) and data on the variability of both species are given. Trogulus tricarintus hirtus Dahl, 1903 is closely related to T. falcipenis, spec. nov., but is distinguishable by long, vertical-standing hairs an the legs. T. falcipenis, spec. nov. is known from Austria (Carinthia), Slovenia, Croatia, Yugoslavia (Montenegro, Serbia) and Albania. Particularly in the northern part of its distribution the species inhabits the litter mainly of beech- and mixed forests and seems to occur mainly in the higher montane-zone. The associated harvestmen-fauna is listed and the endangered status of the species is discussed. Keywords: Trogulus falcipenis, tricarinatus, tricarinatus hirtus, new species, taxonomy, morphometric, variability, aut¬ecology, Trogulidae, Opiliones, Austria, Carinthia, Karawanken, Slovenia, Croatia, Serbia, Montenegro, Albania.
Article
Full-text available
An update of the systematics and determination key of the Opiliones suborder Dyspnoi is provided. The included catalogue represents the first comprehensive species and synonymy listing since Roewer (1923). It summarises all taxonomic changes to date and attempts to be a sound basis against the exponential growing number of online errors, for which examples are given. Species taxonomy features most obvious changes within the Nemastomatidae. The number of species in the collective genus Nemastoma is reduced from 96 (Hallan 2005) to its sensu stricto definition of 7, and the excluded names are transferred to other genera or considered as nomina dubia, predominantly in Paranemastoma. The systematics of the superfamily Ischyropsalidoidea is discussed and family-level diagnoses are renewed to support taxonomical changes: The morphological heterogeneity in the Sabaconidae is resolved by reverting the family to its original monogeneric state. Taracus and Hesperonemastoma are separated as Taracidae fam. n., and Crosbycus is tentatively transferred to this assembly. The remaining genera of Ceratolasmatidae, Acuclavella and Ceratolasma, are included as subfamily Ceratolasmatinae in the Ischyropsalididae and Ischyropsalis is assigned subfamily status, respectively. Other nomenclatural acts are restricted to species-group level with the following synonymies established: Sabacon jonesi Goodnight & Goodnight, 1942 syn. n. (=cavicolens (Packard, 1884)), Dicranolasma diomedeum Kulczyński, 1907 syn. n. (=hirtum Loman, 1894), Mitostoma (Mitostoma) sketi Hadži, 1973a syn. n. (=chrysomelas (Hermann, 1804)), Mitostoma asturicum Roewer, 1951 syn. n. (=pyrenaeum (Simon, 1879a)), Nemastoma formosum Roewer, 1951 syn. n. (=Nemastomella bacillifera bacillifera (Simon, 1879a)), Nemastoma reimoseri Roewer, 1951 syn. n. (=Paranemastoma bicuspidatum (C.L. Koch, 1835)), Nemastoma tunetanum Roewer, 1951 syn. n. (=Paranemastoma bureschi (Roewer, 1926)), Phalangium flavimanum C.L. Koch, 1835 syn. n. (=Paranemastoma quadripunctatum (Perty, 1833)), Crosbycus graecus Giltay, 1932 syn. n. (=Paranemastoma simplex (Giltay, 1932)), Nemastoma bimaculosum Roewer 1951 syn. n. (=Paranemastoma titaniacum (Roewer, 1914)), Trogulocratus tunetanus Roewer, 1950 syn. n. (=Calathocratus africanus (Lucas, 1849)), Trogulus albicerus Sørensen, 1873 syn. n. (=Calathocratus sinuosus (Sørensen, 1873)), Metopoctea exarata Simon, 1879a syn. n. (=Trogulus aquaticus Simon, 1879a), Trogulus galasensis Avram, 1971 syn. n. (=Trogulus nepaeformis (Scopoli, 1763)) and Trogulus roeweri Avram, 1971 syn. n. (=Trogulus nepaeformis (Scopoli, 1763)). Paranemastoma werneri (Kulczyński, 1903) is elevated from subspecies to species. Ischyropsalis luteipes Simon, 1872b is proposed as nomen protectum, taking precedence over Lhermia spinipes Lucas 1866 nomen oblitum. The same accounts for Anelasmocephalus cambridgei (Westwood, 1874) nomen protectum, taking precedence over Trogulus violaceus Gervais, 1844 nomen oblitum, Trogulus closanicus Avram, 1971 nomen protectum over Trogulus asperatus C.L. Koch, 1839a nomen oblitum, as well as Trogulus martensi Chemini, 1983 nomen protectum over Trogulus tuberculatus Canestrini, 1874 nomen oblitum. New combinations, all from Nemastoma, are Histricostoma anatolicum (Roewer, 1962), Mediostoma globuliferum (L. Koch, 1867), Nemastomella hankiewiczii (Kulczyński, 1909), Nemastomella maarebense (Simon, 1913), Nemastomella monchiquense (Kraus, 1961) and Paranemastoma simplex (Giltay, 1932); from Mitostoma: Nemastomella armatissima (Roewer, 1962). Revived combinations are Nemastomella cristinae (Rambla, 1969) (from Nemastoma) and Nemastomella sexmucronatum (Simon, 1911) (from Nemastoma). The following Nemastoma are transferred to Paranemastoma but suggested as nomina dubia: aeginum (Roewer, 1951), amuelleri (Roewer, 1951), bolei (Hadži, 1973a), caporiaccoi (Roewer, 1951), carneluttii (Hadži, 1973a), ferkeri (Roewer, 1951), gigas montenegrinum (Nosek, 1904), gostivarense (Hadži, 1973a), ikarium (Roewer, 1951), quadripunctatum ios (Roewer, 1917), kaestneri (Roewer, 1951), longipalpatum (Roewer, 1951), macedonicum (Hadži, 1973a), multisignatum (Hadži, 1973a), nigrum (Hadži, 1973a), perfugium (Roewer, 1951), santorinum (Roewer, 1951), senussium (Roewer, 1951), sketi (Hadži, 1973a), spinosulum (L. Koch, 1869). Further suggested nomina dubia are Trogulus coreiformis C.L. Koch, 1839a, Trogulus lygaeiformis C.L. Koch, 1839a and Trogulus templetonii Westwood, 1833.
Travaux de l'Institut de Spéologie
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Avram, S., 1971: Quelques espèces nouvelles ou connues du genre Trogulus Latr. (Opiliones). Travaux de l'Institut de Spéologie "Émile Racovitza", 10: 245-272.
Trogulus martensi n. sp. dallʼItalia settentrionale (Arachnida Opiliones)
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Chemini, C., 1983: Trogulus martensi n. sp. dallʼItalia settentrionale (Arachnida Opiliones). Bollettino della Società entomologica italiana, 115(8-10): 125-129.
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Chemini C., 1984. Sulla presenza di Trogulus closanicus AVRAM in Austria, Baviera e Slovenia (Arachnida, Opiliones). Berichte des Naturwissenschaftlich-medizinischen Vereins in Innsbruck, 71: 57-61.
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