© Author(s) 2020. CC Atribution 4.0 License
Revision of species Plagiolophus sulcatus Beurlen, 1939 
(Decapoda, Brachyura) from the Oligocene of Hungary and 
Slovenia 
Revizija vrste Plagiolophus sulcatus Beurlen, 1939 (Decapoda, Brachyura) iz 
oligocena Madžarske in Slovenije 
Matúš HYŽNÝ1, Rok GAŠPARIČ2 & Alfréd DULAI3
1Department of Geology and Palaeontology, Faculty of Natural Sciences, Comenius University, Ilkovičova 6, 
Bratislava 842 15, Slovakia; e-mail: hyzny.matus@gmail.com
2Oertijdmuseum Boxtel, Bosscheweg 80, 5293 WB Boxtel, the Netherlands; e.mail: rok.gasparic@gmail.com
3Department of Palaeontology and Geology, Hungarian Natural History Museum, Ludovika tér 2, Budapest 1088, 
Hungary; e-mail: dulai.alfred@nhmus.hu (Postal address: P.O. Box 137, Budapest 1431) 
Prejeto / Received 14. 1. 2020; Sprejeto / Accepted 9. 4. 2020; Objavljeno na spletu / Published online 22. 4. 2020
Key words: Decapod crustaceans, crab, Glyphithyreus, Oligocene, Rupelian, Chattian 
Ključne besede: raki deseteronožci, rakovica, Glyphithyreus, oligocen, rupelij, chattij
Abstract
The crab species Plagiolophus sulcatus Beurlen, 1939 from the Oligocene (Rupelian) Kiscell Clay of Hungary 
is revised and its holotype is reillustrated for the first time since its original publication. Material from the upper 
Oligocene (Chattian) of Trbovlje (Slovenia) is here considered conspecific with P. sulcatus. Attribution of this 
species to the genus Glyphithyreus, as proposed by Hiroaki Karasawa and Carrie Schweitzer in 2004, is confirmed. 
Glypthithyreus sulcatus differs from congeners in possessing protogastric regions that are subtriangular in 
outline and in having fewer and coarser tubercles on elevated carapace regions.
Izvleček
Revidirana je vrsta rakovice Plagiolophus sulcatus Beurlen, 1939, iz oligocenskih (rupelijskih) kiscellijskih 
glinenih plasti. Prvič po prvotni objavi je predstavljen holotip in novi primerek iz zgornjega oligocena (chattija) 
Trbovelj (Slovenija), ki prav tako pripada vrsti P. sulcatus. Potrjena je pripadnost te vrste rodu Glyphithyreus, 
kar sta predlagala Hiroaki Karasawa in Carrie Schweitzer leta 2004. Glypthithyreus sulcatus se od drugih 
pripadnikov tega rodu razlikuje po tem, da ima trikotno obliko protogastrične regije in manjše število, a 
izrazitejše oblikovane bradavice na višjih delih oklepa.
GEOLOGIJA 63/1, 83-91, Ljubljana 2020
https://doi.org/10.5474/geologija.2020.009
Introduction
Beurlen (1939) described a decapod crusta-
cean faunule from the Oligocene Kiscell Clay of 
Hungary. The ghost shrimps of this assemblage 
have since received proper re-evaluation (Hyžný 
& Dulai, 2014), the three species of brachyu-
ran crabs, including Plagiolophus sulcatus, re-
mained unrevised in respect with modern clas-
sification until now. This species was tentatively 
retained in the genus Plagiolophus Bell, 1858 
(non Pomel, 1857) by Karasawa & Schweitzer 
(2004) in their revision of Glyphithyreus Reuss, 
1859. Those authors noted that, “the placement 
of G. sulcatus is somewhat tentative and is based 
upon our translation of Beurlen’s (1939) original 
description in German and the very poorly repro-
duced illustration in our copy of the work (Kara-
sawa & Schweitzer, 2004, p. 148)“. Thus, since the 
erection of the species by Beurlen (1939), the type 
material of P. sulcatus has not yet been re-exa-
mined.
Bittner (1884) presented an extensive over-
view of Cenozoic sedimentary rocks and their 
fossil contents in the vicinity of Sagor (nowadays 
Zagorje ob Savi) and Trifail (nowadays Trbovlje). 
Among other faunal elements, Bittner (1884: 29) 
also mentioned the presence of a crab that was 
morphologically close to Plagiolophus. Several 
crab specimens from Trbovlje have recently been 
traced by one of us (MH) during a detailed screen 
84 Matúš HYŽNÝ, Rok GAŠPARIČ & Alfréd DULAI
of the main fossil collections in Austria (Hyžný 
& Gross, 2016; Hyžný & Zorn, in press). One of 
these indeed represents Plagiolophus (= Glyph-
ithyreus) and has been considered to be conspe-
cific with P. sulcatus by Hyžný & Gross (2016). 
However, this decision was not based on a first-
hand examination of the type material.
The aim of the present note is to provide a re-
vised description of Glyphithyreus sulcatus on 
the basis of the type specimen of Plagiolophus 
sulcatus from Hungary and of additional materi-
al from Slovenia.
Geological settings
The material that forms the basis for the pres-
ent study comes from two localities, as follows:
Budapest area (Hungary): the holotype of Pla-
giolophus sulcatus originated from the Kiscell 
Clay of Óbuda (currently a part of the city of Bu-
dapest; Fig. 1). The Kiscell Clay Formation con-
sists of grey, well-bioturbated, calcareous clay 
and clayey marl (Báldi, 1983), the type area being 
Óbuda, where brickyards were in operation dur-
ing the second half of the 19th century. The most 
famous of these was the Újlak brickyard (former-
ly Holzspach brickyard); this was the type local-
ity of Plagiolophus sulcatus.
The calcareous nannoflora of the Kiscell Clay 
is indicative of the lower part of zone NP 24 (up-
per Kiscellian) (compare Nagymarosy & Bál-
di-Beke, 1988). This assemblage probably equates 
with the topmost part of zone P 20 and the lower 
part of zone P 21 in the planktonic foraminife-
ral zonation (Horváth, 1998). In the upper part 
of the Kiscell Clay, the assemblage also belongs 
to the upper Kiscellian (NP 24 nannoplankton 
zone and P 21 planktonic foraminiferal zone) (see 
Horváth, 1998, 2002). K-Ar dating of glauconite 
from the Kiscell Clay at Pilisborosjenő, north of 
Budapest, has yielded a date of 33+/-3 Ma (Báldi 
et al., 1975). The Kiscellian is a regional stage in 
the Central Paratethys that is used for part of the 
Lower Oligocene (Rupelian). It was first proposed 
by Báldi (1979) and later defined in a type section 
by Báldi (1986). The Kiscellian is now considered 
to correspond with the Rupelian (Báldi et al., 
1999; Piller et al., 2007).
Generally speaking, the Kiscell Clay is not 
very rich in macrofossils. Strata assigned to this 
unit, however, were mined at several brickyards 
along the margins of the Buda Mountains for 
nearly a century, which explains why their fauna 
is relatively well known, including foraminifera 
(Hantken, 1875; Majzon, 1966; Sztrákos, 1974; 
Fig. 1. Left – Simplified map of Hungary and the Budapest area with the position of the former Újlak brickyard (asterisk). 
Right – Simplified lithostratigraphical scheme of the Hungarian Oligocene at the Buda Hills area (modified after Császár, 
1997); the asterisk indicate approximate position of the Kiscell Clay decapod assemblage. 1 = Hárshegy Sandstone Formation
85Revision of species Plagiolophus sulcatus Beurlen, 1939 (Decapoda, Brachyura) from the Oligocene of Hungary and Slovenia
Gellai-Nagy, 1988; Horváth, 2002, 2003), gas-
tropods and bivalves (Noszky, 1939, 1940; Bál-
di, 1986), cephalopods (Szörényi, 1933; Wagner, 
1938); brachiopods (Meznerics, 1944), ostracods 
(Monostori, 1982, 2004), cirripedes (Szörényi, 
1934), decapod crustaceans (Beurlen, 1939; Hyžný 
& Dulai, 2014), and fishes (Weiler, 1933, 1938; Nolf 
& Brzobohatý, 1994; Szabó & Kocsis, 2016). 
Trbovlje (Slovenia): The locality of Trbovlje 
is situated in the Laško Syncline and belongs to 
geotectonic unit of the Sava folds (Placer, 1999; 
Jelen & Rifelj, 2002). Oligocene and Miocene sed-
imentary rocks were laid down disconformably 
on Triassic and Cretaceous fine-grained, clastic 
rocks (Hafner, 2000). Successive regressive and 
transgressive sequences suggest alternating cy-
cles of deepening and shallowing in the deposi-
tional environment. The stratigraphical sequence 
also shows a variably strong influence of marine 
and terrestrial conditions.
The locality studied is a disused coal pit (GPS 
co-ordinates: 46°08’56” N, 15°04’03” E), situated 
some 3 km east of the city of Trbovlje, along the 
road to Hrastnik (Fig. 2). The area was inten-
sively mined for lignite (brown coal) during the 
last two centuries. On account of the rich brown 
coal deposits, the area has been thoroughly stud-
ied in the past (Bittner, 1884; Petrascheck, 1952; 
Kuščer, 1967; Jelen et al., 1992; Placer, 1999; Haf-
ner, 2000).
The Cenozoic sequence here starts with the 
upper Oligocene Trbovlje Formation, which 
disconformably overlies Triassic rocks. The 
coal-bearing Trbovlje Formation is also known 
as the Socka beds (“Sotzkaschichten”) or Pseu-
do-Socka beds in the older literature (Bechtel et 
al., 2004). This unit starts with basal conglom-
erates, sandstones layers and greyish coloured 
marls to marly limestones. The marly beds con-
tained an economically important coal seam. 
Pollen and coal analysis have demonstrated the 
taxodiacean–cupressacean origin of the main 
coal seam (Bruch, 1998; Križnar, 2000) and most 
likely a transition to a reed marsh in the upper 
part. The overlying marls and marly limestones 
are the most fossil-rich beds (Fig. 2), with diverse 
molluscan and fish assemblages (Križnar, 2015; 
Buckeridge, in press) and abundant floral re-
mains (Lorencon, 2019). The sequence continues 
with a horizon of grey marine clay of the Sivica 
Formation. In the top part of the clay succession 
occur individual layers and lenses of fine-grained 
clastic rocks, particularly sandstones and con-
glomerates. The transition to the clastic beds of 
the lower Miocene Govce Formation is continu-
ous (Hafner, 2000).
The crab-bearing strata of the Trbovlje For-
mation are Late Oligocene in age (Odin et al., 
1994; Bechtel et al., 2004).
Material and methods
The crabs studied herein are part of historical 
collections and have not been prepared further. 
Specimens were photographed with and without 
ammonium chloride coating.
Abbreviations
GBA: Geological Survey of Austria, Vienna 
(Austria).
HNHM: Department of Palaeontology and 
Geology, Hungarian Natural History Museum, 
Budapest (Hungary).
UMJGP: Department for Geology & Palaeon-
tology, Universalmuseum Joanneum, Graz (Aus-
tria).
Fig. 2. Left – Simplified map of Slovenia and locality of provenance (star) of specimen of Glyphithyreus sulcatus (Beurlen, 
1939) studied herein. Right – Simplified lithostratigraphical section of the Trbovlje locality (modified after Bechtel et al., 
2004); strata that have yielded crab specimens are marked.
86 Matúš HYŽNÝ, Rok GAŠPARIČ & Alfréd DULAI
Order Decapoda Latreille, 1802
Infraorder Brachyura Latreille, 1802
Subsection Heterotremata Guinot, 1977
Superfamily Xanthoidea MacLeay, 1838
Family Panopeidae Ortmann, 1893
Subfamily Eucratopsinae Stimpson, 1871
Genus Glyphithyreus Reuss, 1859
(= Plagiolophus Bell, 1858, non Pomel, 1857)
Type species: Glyphithyreus formosus Reuss, 
1859, by original designation.
Diagnosis: See Karasawa & Schweitzer (2004: 
147).
Glyphithyreus sulcatus (Beurlen, 1939) emend.
Figures 3–5
*1939 Plagiolophus sulcatus Beurlen, p. 155, 
pl. 7, fig. 11.
2004 Glyphithyreus sulcatus (Beurlen) – Ka-
rasawa & Schweitzer, p. 148.
2010 Glyphithyreus sulcatus (Beurlen) – 
Schweitzer et al., p. 121.
2016 Glyphithyreus sulcatus (Beurlen) – 
Hyžný & Gross, p. 110, fig. 15.1.
Emended diagnosis: Carapace subhexagonal 
in outline, widest in anterior one-third of length; 
fronto-orbital margin about 65 per cent of max-
imum carapace width; carapace grooves and 
regions well defined, with granular transverse 
ridges; regions covered with coarse granules at 
elevations; protogastric regions subtriangular in 
outline.
Material studied: HNHM M.59.4692, a 
near-complete carapace, the holotype of Plagi-
olophus sulcatus; Óbuda, Hungary (Fig. 3); UM-
JGP 56664, a near-complete individual, retaining 
pereiopods, inclusive of chelipeds, from Trbovlje, 
Slovenia; GBA 2007/024/0005 (Fig. 4A), counter-
part of UMJGP 56664 from Trbovlje, Slovenia 
(Figs. 4B–C). Interestingly, part and counterpart 
of the specimen from Trbovlje ultimately land-
ed up in two collections (see also Hyžný & Gross, 
2016, fig. 15.1; Hyžný & Zorn, in press, pl. 25, fig. 
2).
Description: Carapace subhexagonal in out-
line; L/W (length/width) ratio 0.8, widest in 
anterior one-third of carapace. Fronto-orbital 
margin about 65 per cent of maximum carapace 
width; front broken; orbits poorly preserved. 
Anterolateral margin strongly convex with four 
blunt teeth, including outer orbital tooth; pos-
terolateral margin sinuous, converging posteri-
orly. Carapace grooves and regions well defined; 
epigastric regions well developed, rectangular 
in outline; protogastric regions subtriangular in 
outline, with steep ridges anteriorly; mesogas-
tric region well developed, with elongate, narrow 
anterior process; metagastric region with gran-
ular transverse ridge and two distinct gastric 
pits posteriorly, separated from smooth urogas-
tric region by narrow groove; cardiac region as 
wide as metagastric region, with broad, granular 
transverse ridge; hepatic regions well defined, 
delimited by deep cervical groove posteriorly; 
branchial region divided into two portions by 
distinct branchio-cardiac groove, each bearing 
granular transverse ridge. Regions covered with 
coarse granules at elevations, with cardiac region 
Fig. 3. Glyphithyreus sulcatus (Beurlen, 1939), the holotype of Plagiolophus sulcatus (HNHM M.59.4692) from the Kiscellian 
(Rupelian) of the Budapest area, Hungary. A – Frontal view. B – Dorsal view. C – Left lateral view. D – Ventral view. The spe-
cimen was coated with ammonium chloride prior to photography. Scale bar equals 10 mm.
87Revision of species Plagiolophus sulcatus Beurlen, 1939 (Decapoda, Brachyura) from the Oligocene of Hungary and Slovenia
Fig. 4. Glyphithyreus sulcatus (Beurlen, 1939) from the upper Oligocene (Chattian) of Trbovlje, Slovenia. A – GBA 2007/024/0005 
(unwhitened). B – UMJGP 56664 (unwhitened). C – UMJGP 56664 (whitened with ammonium chloride). Scale bars equal 
10 mm.
being densely granulated, whereas protogastric, 
meso- and metagastric and branchial regions 
having only limited number of relatively large 
tubercles. Chelipeds (pereiopods 1) with robust 
chelae, insufficiently preserved; carpus subquad-
rate in outline; manus approximately two times 
longer than tall, converging proximally; fingers 
shorter than manus. Pereiopods 2–5 slender, dis-
tal elements not preserved sufficiently.
Remarks: Karasawa & Schweitzer (2004, p. 
148) noted that, “the description of G. sulcatus 
clearly indicates two transverse ridges on the 
branchial regions, separated by a very deep cav-
ity, which is certainly characteristic of Glyphi-
thyreus.” We can confirm this and thus corrobo-
rate the transfer of this species to this genus.
As far as carapace outline is concerned, 
Glyphithyreus sulcatus appears to be close to G. 
88 Matúš HYŽNÝ, Rok GAŠPARIČ & Alfréd DULAI
ellipticus Bittner, 1875 from the Eocene of Italy 
(Bittner, 1875), as far as the published figure al-
lows to judge this. However, the latter differs in 
having more rounded protogastric regions; these 
are subtriangular in outline in G. sulcatus. Ad-
ditionally, G. sulcatus has fewer granules on the 
elevated parts of carapace regions (Figs. 3, 5). In 
this respect, this species differs from all conge-
ners known to date, including G. formosus Re-
uss, 1859 and G. wetherellii (Bell, 1858), in which 
carapace regions have a much finer granulation 
distributed over a larger area. Moreover, G. for-
mosus has a wider fronto-orbital margin (Reuss, 
1859, pl. 2, fig. 1) than G. sulcatus.
Conclusions
A revised description of Plagiolophus sulcatus, 
based both on its type specimen from the lower 
Oligocene (Rupelian) of Hungary and additional 
material from the upper Oligocene (Chattian) of 
Trbovlje (Slovenia), is presented. Interestingly, 
part and counterpart of the specimen from Tr-
bovlje were transferred to the Universal Museum 
Joanneum at Graz and the Geological Survey at 
Vienna. The holotype of the species is refigured 
for the first time here since its original publica-
tion. Attribution of P. sulcatus to Glyphithyreus, 
first suggested by Karasawa & Schweitzer (2004), 
is confirmed. Comparison with congeners sug-
gests that G. sulcatus is differentiated by having 
subtriangular protogastric regions and fewer and 
coarser tubercles on elevated carapace regions.
Acknowledgements
Martin Gross (UMJGP) and Irene Zorn (GBA) 
granted access to the material studied. Yusuke 
Ando (Mizunami Fossil Museum, Japan) and John 
W. M. Jagt (Natuurhistorisch Museum Maastricht, 
Maastricht, the Netherlands) commented on an earlier 
version of the typescript. This research was supported 
by the Slovak Research and Development Agency un-
der contract no. APVV-17-0555 and by the Hungarian 
Scientific Research Fund (OTKA/NKFIH K112708).
References
Báldi, T. 1979: Changes of Mediterranean 
(?Indopacific) and Boreal influences in 
Hungarian marine mollusc faunas since 
Kiscellian until Eggenburgian times. The sta-
ge Kiscellian. Annales Géologiques des Pays 
Helléniques, VII. Congress CMNS, 1: 19–49.
Báldi, T. 1983: Magyarországi oligocén és alsó 
miocén formációk [Oligocene and Lower 
Micene formations of Hungary]. Akadémiai 
Kiadó, Budapest: 293 p. (in Hungarian)
Báldi, T. 1986: Mid-Tertiary stratigraphy and 
paleogeographic evolution of Hungary. 
Akadémiai Kiadó, Budapest: 201 p.
Báldi, T., Báldi-Beke, M., Horváth, M., 
Nagymarosy, A., Balogh, K. & Sós, E. 1975: 
On the radiometric age and the biostratigrap-
hic position of the Kiscell Clay in Hungary. 
Proceedings of the VIth Congress of RCMNS, 
Bratislava, Veda: 315–317.
Fig. 5. Glyphithyreus sulca-
tus (Beurlen, 1939), recon-
struction scheme of dorsal 
carapace. Abbreviations: 
eg = epigastric region, msg 
= mesogastric regions, mtg 
= metagastric region, pg = 
protogastric region, ug = 
urogastric region.
89Revision of species Plagiolophus sulcatus Beurlen, 1939 (Decapoda, Brachyura) from the Oligocene of Hungary and Slovenia
Báldi, T., Less, G. & Mandic, O. 1999: Some new 
aspects of the lower boundary of the Egerian 
stage (Oligocene, chronostratigraphic scale 
of the Paratethyan area). Abhandlungen der 
Geologischen Bundesanstalt, 56: 653–668.
Bechtel, A., Markic, M., Sachsenhofer, R.F., Jelen, 
B., Gratzer, R., Lücke, A. & Püttmann, W. 2004: 
Paleoenvironment of the upper Oligocene 
Trbovlje coal seam (Slovenia). International 
Journal of Coal Geology, 57/1: 23–48. https://
doi.org/10.1016/j.coal.2003.08.005
Bell, T. 1858: A monograph of the fossil mala-
costracous Crustacea of Great Britain, Part 
I. Crustacea of the London Clay, 44 p. The 
Palaeontographical Society, London.
Beurlen, K. 1939: Neue Dekapoden-Krebse aus 
dem ungarischen Tertiär. Paläontologische 
Zeitschrift, 21: 135–161.
Bittner, A. 1875: Die Brachyuren des vicentini-
schen Tertiärgebirges. Denkschriften der 
kaiserlichen Akademie der Wissenschaften, 
mathematisch-natur wissenschaft l iche 
Classe, 34: 63–106.
Bittner, A. 1884: Die Tertiär-Ablagerungen von 
Trifail und Sagor. Jahrbuch der kaiserlich-
-königlichen Geologischen Reichsanstalt, 34: 
236–281.
Bruch, A. 1998: Palynologische Untersuchungen 
im Oligozän Sloweniens - Paläo-Umwelt und 
Paläoklima im Ostalpenraum. Tübinger Mi-
kropaläontologische Mitteilungen, 18: 1–193.
Buckeridge, J.S., Kočí, T., Gašparič, R. & Kočová 
Veselská, M. in press: Actinobalanus? slove-
niensis (Thoracica, Balanoidea), a new species 
of cirripede from the Oligocene and Miocene 
of Slovenia that grew attached to wood 
substrates. Neues Jahrbuch für Geologie und 
Paläontologie – Abhandlungen. Accepted in 
press March 9th, 2020
Császár, G. (ed.) 1997: Basic Lithostratigraphic 
Units of Hungary. Charts and Short 
Descriptions. Magyar Állami Földtani 
Intézet, Budapest: 114 p.
Gellai-Nagy, Á. 1988: Delineation of Hantken’s 
foraminiferal species from the original co-
llection. A Magyar Állami Földtani Intézet 
Évi Jelentése 1988 évről: 133–173.
Guinot, D. 1977: Propositions pour une nouvel-
le classification des Crustacés Décapodes 
Brachyoures. Comptes rendus hebdomadaires 
des séances de l’Académie des sciences, D285: 
1049–1052.
Hafner, G. 2000: Sedimentološke značilnosti tr-
boveljskih plasti. Unpublished MSc thesis, 
University of Ljubljana, Ljubljana: 105 p.
Hantken, M. 1875: A Clavulina Szabói rétegek 
faunája, I. rész. Foraminiferák (Die Fauna 
der Clavulina Szabói Schichten, I. Teil. 
Foraminiferen). Jahrbuch der königlichen 
ungarischen geologischen Anstalt, 4: 1–93.
Horváth, M. 1998: Paleobathymetrical analysis of 
Upper Eocene-Lower Miocene Foraminifera 
of the Hungarian Paleogene Basin. Acta 
Geologica Hungarica, 41/2: 223–262.
Horváth, M. 2002: Data to revision and distribu-
tion of small foraminifera species described 
by Hantken (1868, 1875). Part I, Textulariidae 
and Miliolidae. Fragmenta Palaeontologica 
Hungarica, 20: 25–42.
Horváth, M. 2003: Data to revision and dis-
tribution of small Foraminifera species 
described by Hantken (1868, 1875). Part II. 
Nodosariidae and Vaginulinidae. Fragmenta 
Palaeontologica Hungarica, 21: 5–32.
Hyžný, M. & Dulai, A. 2014: Deep-water fosso-
rial shrimps from the Oligocene Kiscell Clay 
of Hungary: taxonomy and palaeoecology. 
Acta Palaeontologica Polonica, 59/4: 947–965. 
https://doi.org/10.4202/app.2012.0078
Hyžný, M. & Gross, M. 2016: From the palaeon-
tological collection of the Universalmuseum 
Joanneum – The Cenozoic decapod crusta-
ceans (Crustacea: Malacostraca: Decapoda). 
Joannea, Geologie und Paläontologie, 12: 
73–127.
Hyžný, M. & Zorn, I. in press: A catalogue of 
the fossil decapod crustaceans in the col-
lections of the Geological Survey of Austria 
in Vienna. Abhandlungen der Geologische 
Bundesanstalt.
Jelen, B. & Rifelj, H. 2002: Stratigraphic struc-
ture of the B1 Tertiary tectonostratigraphic 
unit in eastern Slovenia. Geologija, 45/1: 115–
138. https://doi.org/10.5474/geologija.2002.010
Jelen, B., Aničič, B., Brezigar, A., Buser, S., 
Cimerman, F., Drobne, K., Monostori, M., 
Kedves, M., Pavšič, J. & Skaberne, D. 1992: 
Model of positional relationships for Upper 
Paleogene and Miocene strata in Slovenia. 
Interdisciplinary Geological Conference 
on the Miocene Epoch, Abstracts and Field 
Trips. Ancona, 1–72.
Karasawa, H. & Schweitzer, C.E. 2004: 
Revision of the genus Glyphithyreus Reuss, 
1859 (Crustacea, Decapoda, Brachyura, 
Xanthoidea) and recognition of a new genus. 
Paleontological Research, 8: 143–154.
Križnar, M. 2000: Ksilotomske analize rjavih 
premogov iz Zasavja. Unpublished BSc the-
sis, University of Ljubljana, Ljubljana: 62 p.
90 Matúš HYŽNÝ, Rok GAŠPARIČ & Alfréd DULAI
Križnar, M. 2015: Oligocenska ihtiofavna 
(Teleostei) Slovenije - pregled in problematika 
najdišč. In: Rožič, B. (ed.): Razprave, poroči-
la = Treatises, reports / 22. posvetovanje slo-
venskih geologov = 22nd Meeting of Slovenian 
Geologists. Geološki zbornik, 23: 99–104.
Kuščer, D. 1967: Zagorski terciar (Tertiary for-
mations of Zagorje). Geologija, 10: 5–85.
Latreille, P.A. 1802: Histoire naturelle, générale 
et particulière des Crustacés et des Insectes. 
Ouvrage faisant suite à l’histoire naturelle 
générale et particulière, composée par Leclerc 
de Buffon, et rédigée par C.S. Sonnini, mem-
bre de plusieurs sociétés savantes. Familles 
naturelles des genres. Vol. 3. F. DuFart, Paris: 
467 p.
Lorencon, R. 2019: Oligocenska flora v okolici 
Trbovelj. Konkrecija, 8: 12–18.
MacLeay, W.S. 1838: On the brachyurous deca-
pod Crustacea brought from the Cape by Dr. 
Smith. In: Smith, A. (ed.) Illustrations of the 
Annulosa of South Africa; being a portion 
of the objects of natural history chiefly col-
lected during an expedition into the interi-
or of South Africa, under the directin of Dr. 
Andrew Smith, in the years 1834, 1835. and 
1836; fitted out by “The Cape of Good Hope 
Association for Exploring Central Africa”. 
Smith, Elder, and Co., London: 53–71.
Majzon, L. 1966: Foraminifera-vizsgálatok. 
Akadémiai kiadó, Budapest: 939 p.
Meznerics, I. 1944: Die Brachiopoden des unga-
rischen Tertiärs. Annales historico-naturales 
Musei nationalis hungarici, 36: 10–60.
Monostori, M. 1982. Oligocene ostracods from 
the surroundings of Budapest. Annales 
Universitatis Scientiarum Budapestinensis de 
Rolando Eötvös Nominatae, Sectio Geologica, 
21: 31–102.
Monostori, M. 2004: Lower Oligocene (Kiscellian) 
ostracods in Hungary. Annales Universitatis 
Scientiarium Budapestinensis, Sectio 
Geologica, 34: 27–141.
Nagymarosy, A. & Báldi-Beke, M. 1988: The 
position of the Paleogene formations of 
Hungary in the standard nannoplankton zo-
nation. Annales Universitatis Scientiarum 
Budapestinensis de Rolando Eötvös 
Nominatae, Sectio Geologica, 28: 3–25.
Nolf, D. & Brzobohatý, R. 1994: Fish otoliths from 
the Late Oligocene (Eger and Kiscell formati-
ons) in the Eger area (northeastern Hungary). 
Bulletin de l’Institut royal des Sciences natu-
relles de Belgique, Sciences de la Terre, 64: 
225–252.
Noszky, J. 1939: A kiscelli agyag mollusz-
ka-faunája. I. rész. Lamellibranchiata 
(Die Mollusken fauna des Kisceller Tones 
(Rupelien) aus der Umgebung von Budapest. 
I. Teil. Lamellibranchiata). Annales histori-
co-naturales Musei nationalis hungarici, 32: 
19–146.
Noszky, J. 1940: A kiscelli agyag molluszka-
faunája. II. rész. Loricata, Gastropoda, 
Scaphopoda. (Die Mollusken fauna des 
Kisceller Tones (Rupelien) aus der Umgebung 
von Budapest. II. Teil. Loricata, Gastropoda, 
Scaphopoda). Annales historico-naturales 
Musei nationalis hungarici, 33: 1–80.
Odin, G.S., Jelen, B., Drobne, K., Uhan, J., 
Skaberne, D., Pavsic, J., Cimerman, F., 
Cosca, M. & Hunziker, J.C. 1994: Premiers 
âges géochronologiques de niveaux volca-
noclastiques oligocènes de la région de Zasavje, 
Slovénie. Giornale di Geologia, 56/1: 199–212.
Ortmann, A.E. 1893: Die Decapoden-Krebse 
des Strassburger Museums, mit besonderer 
Berücksichtigung der von Herrn Dr. Döderlein 
bei Japan und bei den Liu-Kiu-Inseln ge-
sammelten und zur Zeit im Strassburger 
Museum aufbewahrten Formen. VII. Theil. 
Abtheilung: Brachyura (Brachyura genui-
na Boas) II. Unterabtheilung: Cancroidea, 2. 
Section: Cancrinea, 1. Gruppe: Cyclometopa. 
Zoologische Jahrbücher. Abteilung für 
Systematik, Geographie und Biologie der 
Thiere, 7/3: 411–495.
Petrascheck, W. 1952: Der Einfluß der Fazies der 
Flözablagerung auf die Eigenschaften der 
Kohle. Zeitschrift der deutschen geologi-
schen Gesellschaft, 104: 1–9. 
Piller, W.E., Harzhauser, M. & Mandic, O. 2007: 
Miocene Central Paratethys stratigraphy 
– current status and future directions. 
Stratigraphy, 4: 71–88.
Placer, L. 1999: Structural meaning of the Sava 
folds. Geologija, 41: 191–221. https://doi.
org/10.5474/geologija.1998.012
Pomel, A. 1857: Note critique sur les caractères et 
les limites du genre Palaeotherium. Archives 
des Sciences physiques et naturelles Genève, 
5: 200–207.
Reuss, A.E. 1859: Zur Kenntnis fossiler Krabben. 
Akademie der Wissenschaften Wien, 
Denkschrift, 17: 1–90, pls. 1–24.
Schweitzer, C.E., Feldmann, R.M., Garassino, 
A., Karasawa, H. & Schweigert, G. 2010: 
Systematic list of fossil decapod crustacean 
species. Crustaceana Monographs, 10: 1–222. 
https://doi.org/10.1163/ej.9789004178915.i-222
91Revision of species Plagiolophus sulcatus Beurlen, 1939 (Decapoda, Brachyura) from the Oligocene of Hungary and Slovenia
Stimpson, W. 1871: Preliminary report on the 
Crustacea dredged in the Gulf Stream in the 
Straits of Florida by L.F. de Pourtales, Assist. 
U. S. Coast Survey. Part I. Brachyura. Bulletin 
of the Museum of Comparative Zoology at 
Harvard College, 2: 109–160.
Szabó, M. & Kocsis, L. 2016: A preliminary report 
on the Early Oligocene (Rupelian, Kiscellian) 
selachians from the Kiscell formation (Buda 
Mts, Hungary), with the re-discovery of 
Wilhelm Weiler’s shark teeth. Fragmenta 
Palaeontologica Hungarica, 33: 31–64.
Szörényi, E. 1933: Adatok a harmadkori Sepia-
félék ismeretéhez, néhány új magyarorszá-
gi faj alapján. (Neue tertiäre Sepiinae aus 
Ungarn nebst Bemerkungen zum zeitlichen 
Auftreten und zur Entwicklung der Gattung 
Sepia). Földtani Közlöny, 63: 1183–1189.
Szörényi, E. 1934: Oligocén Scalpellum ma-
radványok Magyarországról. (Scalpellum 
Reste aus dem ungarischen Oligozän). 
Földtani Közlöny, 64: 272–277.
Sztrákos, K. 1974: Paleogene planktonic fora-
miniferal zones in northeastern Hungary. 
Fragmenta Mineralogica et Palaeontologica, 
5: 29–81.
Wagner, J. 1938: A kiscelli közép-oligo-
cén (Rupélien) rétegek kétkopoltyús 
Cephalopodái és új Sepia-félék a magyar 
eocénból. (Die dibranchiaten Cephalopoden 
der mitteloligozänen (Rupelien) Tonschichten 
von Kiscell und eine Sepiinae aus dem unga-
rischen Eozän). Annales historico-naturales 
Musei nationalis hungarici, 31: 179–199.
Weiler, W. 1933: Két magyarországi oligocén-
korú halfauna. (Zwei oligozäne Fischfaunen 
aus dem Königreich Ungarn). Geologica 
Hungarica, Series Palaeontologica, 11: 1–54.
Weiler, W. 1938: Neue Untersuchungen an mi-
tteloligozänen Fischen Ungarns. Geologica 
Hungarica, Series Palaeontologica, 15: 1–31.