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 n a pač n o vrednotene vrste  
suhih juž i n z obronkov juž n i h Alp  
(Opiliones: Trogulidae)
Tone NOVA K
1
, Axel L. SCHÖNHOFER
2
, Ljuba SLANA NOVAK
3
,  
Peter KOZEL
1,4
, Saška LIPOVŠEK
1,5,6
, Jochen MARTENS
7,8
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 
missing a thorough reconsideration of the first described species Trogulus nepaeformis Scopoli, 
1763. At the same time, this is 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 
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 NOV AK, Axel L. SCHÖNHOFER, Ljuba SLANA NOV AK, 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
 V taksonomskem in sistematskem smislu je rod suhih južin plošček, Trogulus Latreille, 1802, 
med najbolj zapletenimi taksoni pajkovcev v Evropi. V zadnjih desetletjih so bile razrešene 
mnoge težave in opisane številne nove vrste ploščkov. Kljub temu raznolikost znotraj nekaterih 
skupin še vedno ni zadovoljivo razjasnjena. Med temi so srednje veliki ploščki iz skupine 
nepaeformis, deloma zato, ker prva opisana vrsta iz skupine, Trogulus nepaeformis Scopoli, 
1763, še vedno ni podrobno proučena in opisana v skladu z modernimi standardi. Obenem je 
to prva vrsta suhih južin, opisana z ozemlja Slovenije. Tu podajamo ponovni opis Scopolijeve 
vrste z namenom, da pripravimo verodostojno osnovo za revizijo celotne skupine nepaeformis. 
Za T. nepaeformis je značilno kratko, čokato šilo glans penis, in grozdaste semenske vrečke iz 
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, Kofiniotis Roewer, 1940 and Trogulus Latreille, 1802 ( Martens 
1978; Schönhofer 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 
(Schönhofer 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 
secretion, which is probably released by diffusion through areas of thin cuticle, and glues substrate 
particles onto the surface, which hardens into a soil-encrusted, insoluble mail ( Wolff et al. 2016). 
For study purposes, soil encrustations may be removed by dabbing (not brushing) with a stiff brush 
(Schönhofer & Martens 2009), a time-consuming procedure, which sometimes causes damage 
to fine structures, such as bristles on legs, important for identification of some taxa. All this makes 
morphological identification rather difficult, especially within the most diverse Trogulus lineage 
B sensu Schönhofer & Martens (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 
(Schönhofer & Martens 2010). To complicate matters, two or three medium-sized troguli species 
co-occur in many localities (e.g., Chemini 1983, 1984). Thus, although the taxonomic position of 
Trogulus in Central Europe seems straightforward, it is actually extremely complex ( Martens 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 Schönhofer & Martens (2010). Ta xonom ic a l 
problems within these groups have partly been dealt with morphologically by Latreille (1802), 
Koch (1839), Dahl (1903), Avram (1971), Martens (1978), Weiss (1978), Neuffer (1980), 
Chemini (1983, 1984), Chemini & Martens (1988) and Weiss 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 Avram, 
1971, T. martensi Chemini, 1983 and T. cisalpinus Chemini & Martens, 1988. 
Combining morphological and genetic information provided a progress in Trogulus taxonomy 
in the last two decennia, and revealed a much-underestimated cryptic diversity within the 
genus ( Schönhofer 2008; Schönhofer & Martens 2010). This approach allowed resolving 
a number of open taxonomic and distributional problems ( Schönhofer 2008; Schönhofer & 
Martens 2008, 2009, 2010; Schönhofer 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, Scopoli (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 ( Scopo li, 1 7 6 3 ) . Later , Scopo li ʼ s co llection 
burned in his private house (Guglia 1972; Petkovšek 1977; Bavdaž 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 te xtual descri p ti o n. A t 
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 Martens (1978) and Chemini 
(1983, 1984). Apparently, misinterpretation of Scopoli’s intention often occurred (e.g., Avram 1971, 
Martens 1978, Schönhofer 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 NOV AK, Axel L. SCHÖNHOFER, Ljuba SLANA NOV AK, 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 first one described within a group of related forms. The redescription of the nominal species is 
crucial to resolve the central European Trogulus taxonomy ( Chemini & Martens 1988). 
In this contribution we aim 1) to establish the terra typica and locus typicus of T. nepaeformis, 
2) to identify the most probable candidate for Scopoliʼs species among syntopic, morphologically 
similar, medium-sized troguli, and 3) to redescribe this species to provide a clear view on this 
crucial species and to avoid further identification confusion within the group.
Materials and Methods
In searching for the most probable species described by Scopoli, we first analysed the 
original description of Acarus nepeformis Scopoli, 1763 ( Scopoli 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 (Guglia 1972; Petkovšek 
1977; Wraber 1978; Baker 1999; Bavdaž 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 (Kordiš 1986; Marinček et al. 
1987; Dakskobler 1998; Bavdaž et al. 1999; Gorkič & Cernetič-Gregorič 2000). A further 
relevant cue is the measure used by Scopoli in measurements of his specimens (Scopoli 1763; 
Rottleuthner 1985; Baker 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 
to adjust best visibility of surface structures and bristles. Digital images captured at different 
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 ( Martens 1978, Schönhofer 2008, 
Schönhofer & Martens 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
Identification of the locus typicus 
Scopoli (1763) wrote his volume “Entomologia carniolica” while residing in Idrija (1754–
1769), western Slovenia (in older literature Idria in the Duchy of Carniola) (K ryštufek 2023). 
Despite his collecting journeys to many places in Carniola ( Guglia 1972), he putatively 
described most species from Idrija and its close vicinity (Guglia 1972; Petkovšek 1977; 
Wraber 1978; Baker 1999; Bavdaž 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. 
Identification of most credible candidate species
In 206 collection efforts in 46 localities in Idrija and its close vicinity, the following 
harvestman species have been recorded (own unpublished data):
CYPHOPHTHALMI
Sironidae
Cyphophthalmus duricorius Joseph, 1868
(PALPATORES)
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 Šilhavý, 1938
Opilio parietinus (De Geer, 1778)
Phalangium opilio Linnaeus, 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 NOV AK, Axel L. SCHÖNHOFER, Ljuba SLANA NOV AK, 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 (Hermann, 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 (Scopoli, 1763)
Trogulus oltenicus Avram, 1971
Trogulus nepaeformis sensu Chemini 1983, 1984
Trogulus sp. gr. tricarinatus Linnaeus, 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 Chemini (1983, 1984) belong to the nepaeformis 
species-group (cf. Schönhofer 2008; Schönhofer & Martens 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 sensu Chemini ≈ 
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.
Scopoliʼs (1763: 390) original description is as follows: “1070. ACARUS 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 r o o t s . O b l o n g , l e a f - l ik e d e p r e s s e d, s o i l -
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, Scopoli (1763; Explicatio) used the Paris line for size indication, showing 
lines in three Paris twelfths – linea; tres unciae Parisinae ( Baker 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 ( Rottleuthner 
1985: Küstenland, Krain, Istrien, Friaul). The printed Paris line in Scopoli (1763) measured 2.24 mm 
(Baker 1999), while the Vienna line measured 2.195155 mm, and the Southern line 2.063305 mm 
(Rottleuthner 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, 1 758), Naucoridae ( Gogala et al. 1994), in Scopoli (1763) quoted sub 
Nepa, refers to the anterior portion of this water bug and the hood of troguli; although the shape 
and colour comply with all medium-sized troguli, it best fits with the same species. Scopoli used 
the name antennae for the hood horns, and likely interpreted the space below the hood with 
chelicerae and pedipalps as the “mouth".
In his descriptions, Scopoli cited only few specific localities, which were interpreted by 
Petkovšek (1977), Wraber (1978), Baker (1999) and Bavdaž et al. (1999). Scopoliʼs notation 
that T. nepaeformis had been found several times in moss between beech roots is a case of 
unspecific location citation (cf. Baker 1999). Nevertheless, it is obvious that Scopoli inspected 
more localities close to his living place. In his time, Scopoliʼs house in Idrija was very probably 
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 is about 300 m away from Scopoliʼs 
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 18
th
 and 19
th
 centuries, 
while moderate steep slopes were already used as meadows and pastures in the 18
th
 century 
(M. Lužnik, personal comm.). Afterwards, great care was implied for forest reconstitution. 
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 ( Kordiš 1986; 
Dakskobler 1998; Gorkič & Cernetič-Gregorič 2000). In this respect, the area is in 
harmony with Scopoli’s microhabitat description.
For all the above consideration: The perfectly fitting body size, its abundance greatly 
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) ( Scopoli 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 Schönhofer 
(2008) and Schönhofer & Martens (2010) with ~ roundly shaped head cap (hood), small eyes 
Tone NOV AK, Axel L. SCHÖNHOFER, Ljuba SLANA NOV AK, 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
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 
vesicles; Metatarsus II: Tarsus II lengths in ♂ ≈ 1: 1, in ♀ ≈ 1.15: 1. 
Remark: Until the detailed revision of related medium-sized troguli is provided, this diagnosis 
should be considered contemporary.
Material examined 
Neotype 
SLOVENIA • ♂; Pringl, Idrija; 45.99753° N, 14.01126° E; 539 m a.s.l.; mixed forest, 10 Jun. 
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 
Novak identification code).
Further material of the species considered 
SLOVENIA • Brusovše, Idrija; 1 ♂, 1 ♀, 1 juv.; 46.00° N, 14.03° E; 339 m a.s.l.; Jun. 1997; 
M. Vončina leg. (TN 1712/1998). – 8 ♂♂, 12 ♀♀, 4 juv.; Pringl, Idrija; 46.00° N, 14.01° E; 
539 m a.s.l.; 10 Jun. 2000; L. Slana Novak and T. Novak leg. (TN 234/2000). – Ibid.; 1 ♂, 2 ♀♀, 
2 juv.; 10 Jun. 2000; L. Slana Novak and T. Novak leg. (TN 117/2000). – Ibid.; 1 ♂; 01 Aug. 
2000; T. Novak leg. (TN 384/2000). – Ibid.; 2 ♂♂, 6 ♀♀, 2 juv.; 14 Jul. 2001; B. Bertoncelj and 
T. Novak leg. (TN 226/2001). – Idrija; 19 ♂♂, 65 ♀♀, 32 juv.; 46.00° N, 14.01° E; 441 m a.s.l.; 
B. Bertoncelj and T. Novak leg. (TN 237/2001). Spodnja Idrija; 5 ♂♂, 2 ♀♀, 7 juv.; 46.03° N, 
14.04° E; 570 m a.s.l.; 10 Jun. 2000; L. Slana Novak and T. Novak leg. (TN 145/2000). – Marija 
Snežna, Avče; 1 ♂, 3 ♀♀, 1 juv.; 46.10° N, 13.69° E, 311 m a.s.l.; 02 Aug. 2000; T. Novak leg. 
(292/2000). All PMSL. – Idrija; 4 ♂♂; 440 m a.s.l., 46.00° N, 14.03° E; 31 May 2004; L. Slana 
Novak and T. Novak leg. (Coll. J. Martens CJM 3832 in SMF = Senckenberg Forschungsinstitut 
und Naturmuseum, Frankfurt am Main, Germany); 9 ♀♀, dito CJM 3833).
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 
distance 0.44; body with dense, relatively large, flat, dome-shaped tubercles (smooth papillae) 
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 
fingers; basal article with 3 short, rigid bristles antero-apically, 2–3 such bristles on distal article 
apically, and row of few bristles medio-ventrally; basal article and fingers brown, distal article 
dark beige, mottled with brown spots, fingers with black base and black-tipped. Ch basal article 
length 1.42, distal article length 1.27, movable finger length 0.51.
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 
younger individuals, Ta brownish; except Ta articles with scattered flat dome-shaped tubercles, 
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 
II: Ta II lengths ≈ 1: 1. Cx II pro-medial interdistance: 2.32, Cx II pro-lateral interdistance: 3.10, 
Cx IV retro-medial interdistance: 2.86, Cx IV retro-lateral interdistance: 3.68.
Female (n=1)
Body 8.66 long, 3.51 wide; as in ♂.
Chelicerae as in ♂. Ch basal article length 1.58, distal article length 1.45, movable finger 
length 0.57. 
Pedipalps as in ♂. Article lengths in Table 1. 
Ovipositor 1.72 long, Rec sem acinous, consisted of eight balloon-shaped and one short-
tubular vesicles. 
Legs. As in ♂. Article lengths in Table 1. Mt II: Ta II lengths ≈ 1.15: 1. 
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 Total length
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 NOV AK, Axel L. SCHÖNHOFER, Ljuba SLANA NOV AK, Peter KOZEL, Saška LIPOVŠEK, Jochen MARTENS:  
Redescription of Trogulus nepaeformis (Scopoli, 1763) ... / Ponovni opis Trogulus nepaeformis (Scopoli, 1763) ...

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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.

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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 NOV AK, Axel L. SCHÖNHOFER, Ljuba SLANA NOV AK, Peter KOZEL, Saška LIPOVŠEK, Jochen MARTENS:  
Redescription of Trogulus nepaeformis (Scopoli, 1763) ... / Ponovni opis Trogulus nepaeformis (Scopoli, 1763) ...

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Remarks on taxonomy
Acarus Nepeformis Scopoli, 1763 was placed in the correct arachnid order, Opiliones, by 
Latreille (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) 
(Schönhofer 2013). Trogulus nepaeformis s. s. is very similar to an until now unresolved 
number of additional species belonging to the nepaeformis species-group sensu Schönhofer 
(2008) and Schönhofer & Martens (2010), which have constantly been cited under this name. 
The revision of the group is currently at the very beginning. So far, only few authors authentically 
refer to T. nepaeformis s. s. in the territory of Slovenia (Scopoli 1763; Mršić 1997; Novak & 
Gruber 2000; Tarman 2003; Blick & Komposch 2004; Novak et al. 2006, 2007; Klenovšek 
et al. 2013a, b; Lipovšek et al. 2021). Although body length and width are considerably 
variable, on the one hand, the genital morphology (penis, receptacula seminis) and measures 
of metatarsus II (Table 2) proved to be very constant allowing unambiguous identification of 
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 
(Schönhofer & Martens 2008; Schönhofer 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)
7.39±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 
(0.94–1.03)
♀ 
(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 about hundred specimens, only a few additional localities in Slovenia have been confirmed 
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 
moss, rock fissures, etc. This species seems to prefer mull humus stratum in soils on limestone 
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 ( Pabst 1953, sub T. nepaeformis).

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Discussion
Sharing the Alpine, central European, Dinaric, Sub-Mediterranean and Sub-Pannonian 
biota, most harvestmen including all troguli species in Slovenia belong to the first three 
biogeographical regions. There are more than ten Trogulus species in the country (Hadži 1931; 
Chemini 1984; Komposch 2000; Novak & Gruber 2000; Novak et al. 2006; Schönhofer 
2009; Schönhofer & Martens 2009; Schönhofer & Novak 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 
(Schönhofer 2009). It is important that species of the nepaeformis species-group are similar in 
size and external morphology, but differ in genital morphology and are allopatric in most cases 
(Schönhofer 2009). Moreover, it is encouraging that comparable genetic distances coincide 
with morphospecies (Schönhofer 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 
easier and most probably denotes the correct species. The designation of this neotype satisfies 
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. 
Marta Lužnik provided ample valuable information on the beech forests in the vicinity of 
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 
is co-financed by the Republic of Slovenia, Ministry of Education, Science and Sport and the 
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 
University for field research into Alpine arachnids.
Tone NOV AK, Axel L. SCHÖNHOFER, Ljuba SLANA NOV AK, Peter KOZEL, Saška LIPOVŠEK, Jochen MARTENS:  
Redescription of Trogulus nepaeformis (Scopoli, 1763) ... / Ponovni opis Trogulus nepaeformis (Scopoli, 1763) ...

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