© Author(s) 2020. CC Atribution 4.0 License
On the occurrence of Iphiculus eliasi Hyžný & Gross, 2016 
(Decapoda, Brachyura, Leucosioidea) from the Miocene of 
Catalonia (northeastern Iberian Peninsula) 
Novi podatki o razširjenosti vrste Iphiculus eliasi Hyžný & Gross, 2016 (Decapoda, 
Brachyura, Leucosioidea) iz miocena Katalonije (severovzhod Iberijskega polotoka) 
Àlex OSSÓ1, Matúš HYŽNÝ2, Manu GÓMEZ3, David ALBALAT4 & Fernando A. FERRATGES5
1Llorenç de Villalonga, 17B, 1-1 43007 Tarragona, Catalonia; Museu de Ciències Naturals de Barcelona,  
Paleontologia (Barcelona, Catalonia); e-mail: aosso@comt.cat
2Department of Geology and Palaeontology, Faculty of Natural Sciences, Comenius University, Ilkovičova 6, 
Bratislava, Slovakia. Geological-palaeontological Department, Natural History Museum, Vienna, Austria;  
e-mail: hyzny.matus@gmail.com
3Sant Pere 7, 2n 08720 Vilafranca del Penedès (Catalonia); e-mail: mnu.gomez@gmail.com
4Benvingut Socies, 75, 2n, 43700, el Vendrell, Tarragona; e-mail: dalbalat@colgeocat.org
5Departamento de Ciencias de la Tierra-IUCA, Universidad de Zaragoza, Zaragoza E-50009, Spain;  
e-mail: Ferratges@unizar.es
Prejeto / Received 15. 11. 2019; Sprejeto / Accepted 7. 4. 2020; Objavljeno na spletu / Published online 22. 4. 2020
Key words: Leucosioidea, Iphiculidae, Miocene, Langhian, Catalonia 
Ključne besede: Leucosioidea, Iphiculidae, miocen, langhij, Katalonija
Abstract
Recovery of two specimens of leucosioid crabs in Langhian (middle Miocene) strata at Vilafranca del Penedès 
(Alt Penedès, Catalonia) and a re-examination of another leucosioid from the palaeontological collections of the 
Vinseum (Vilafranca del Penedès, Catalonia) have led us to consider all of these as conspecific with Iphiculus 
eliasi Hyžný & Gross, 2016, described first from the middle Miocene of Austria. The sternal and pleonal remains 
preserved in one of the Catalonian specimens allows to emend the original description of the species. Likewise, 
a specimen from the middle Miocene of Portugal, previously described as a paguroid, is herein transferred to 
this species. The occurrence of I. eliasi, either in outcrops along the northeastern and southwestern coasts of the 
Iberian Peninsula, corroborates the close relationship between decapod assemblages which inhabited similar 
palaeoenvironments in the Central Paratethys, the western Mediterranean and even the nearest Atlantic waters, 
during the middle Miocene.
Izvleček
Dva nova primerka leukosioidnih rakov langhijske (srednjemiocenske) starosti iz kraja Vilafranca del 
Penedès (Alt Penedès, Katalonija) in dodaten primerek iz muzeja Vinseum (Vilafranca del Penedès, Katalonija) 
smo določili kot Iphiculus eliasi Hyžný & Gross, 2016, ki je bil prvič opisan iz srednjemiocenskih plasti Avstrije. 
Na podlagi dobro ohranjenih morfoloških podrobnosti sternuma in pleona lahko dopolnimo originalni opis vrste. 
Primerek iz srednjega miocena Portugalske je določen kot I. eliasi, ki je bil opisan kot ostanek raka samotarja. 
Razširjenost vrste I. eliasi na iberijskem polotoku dodatno potrjuje podobnosti med fosilnimi združbami 
deseteronožcev, ki so v srednjem miocenu poseljevale centralno Paratetido, zahodno Sredozemlje in plitvovodna 
območja v Atlantskem oceanu.
GEOLOGIJA 63/1, 101-108, Ljubljana 2020
https://doi.org/10.5474/geologija.2020.011
Introduction
Miocene decapod crustacean assemblages of 
the Vallès-Penedès and Camp de Tarragona ba-
sins (northeastern Iberian Peninsula) have been 
studied by a number of scholars and are well 
known (Almera, 1896; Via, 1932; Solé & Via, 1989; 
Müller, 1993; Artal, 2008; Garassino et al., 2009; 
Ossó, 2010). Moreover, due to collecting efforts by 
enthusiastic fossil hunters new occurrences are 
constantly being reported, thus expanding our 
knowledge of fossil decapod crustacean assem-
blages of these areas. One such occurrence is re-
corded in the present contribution.
102 Àlex OSSÓ, Matúš HYŽNÝ, Manu GÓMEZ, David ALBALAT & Fernando A. FERRATGES
Müller (1993) summarised Neogene decapod 
crustaceans known at that time from Catalonia 
and described a number of new taxa, mainly 
from the reef limestones of Olèrdola, as well as 
from Vilafranca del Penedès and Santa Margari-
da i Els Monjos (Alt Penedès). More than a decade 
ago, a large number of fossil decapods, mainly 
Palaeopinnixa mytilicola Vía, 1966 were recov-
ered in the so-called Vilafranca marls (Langhi-
an), extracted during construction works for the 
high-speed railway line on the outskirts of Vila-
franca del Penedès. Among these, remains of a 
carapace and a counterpart of a male venter of 
a leucosioid crab were recovered. The taxonom-
ic assessment of this leucosioid is the goal of the 
present report.
Geological setting
The material studied comes from the localities 
of Vilafranca del Penedès and Santa Margarida i 
els Monjos, both within the Vallès-Penedès Basin 
and exposing Miocene strata. This basin repre-
sents a NE-SW-oriented depression limited to the 
northwest and to the southeast by the Prelitoral 
and Litoral ranges, respectively, which are made 
of Palaeozoic and Mesozoic rocks. The Vallès-
Penedès Basin corresponds to the emerged part 
of the NE-SW and NNW-SSE horst and half-gra-
ben system formed during the Oligocene-Mio-
cene opening of the western Mediterranean (Bar-
trina et al., 1992; Cabrera & Calvet, 1996; Roca 
et al., 1999; Cabrera et al., 2004) (Fig. 1). Rifting 
and thermal subsidence related with this open-
ing led to the accumulation of marine and con-
tinental sediments in the Vallès-Penedès Basin, 
Fig. 1. Simplified geological map of the Vallès-Penedès Basin (modified from Roca et al., 2004). The main upper Burdigalian-
Langhian coralgal and calcarenitic-terrigenous facies are indicated. The red stars indicate the location of the outcrops.
103On the occurrence of Iphiculus eliasi Hyžný & Gross, 2016 (Decapoda, Brachyura, Leucosioidea) from the Miocene of...
from the early to late Miocene (early Burdigalian 
to Tortonian), as discussed in detail by Cabrera 
et al. (2004) and Casanovas-Vilar et al. (2016). Al-
though most of the Vallès-Penedès sedimentary 
infill is of terrestrial origin, three transgressions 
occurred during the late Burdigalian, Langhian 
and early Serravallian, leading to the deposition 
of various marine facies (Cabrera et al., 1991; 
Cabrera & Calvet, 1996; Roca et al., 1999). During 
the most significant Langhian transgression, a 
shallow sea developed in the Penedès area, where 
fringing reefs, carbonate platform and ramp 
sediments, open marine marls and transitional 
shales and sands were laid down.
At the Vilafranca site, grey marls with inter-
bedded levels of fine sands, located predominant-
ly at the top of the unit, are exposed. Based on 
borehole data, the unit attains a thickness of ap-
proximately 300 m (Permanyer, 1982; Cabrera et 
al., 1991) and it extends across the Penedès Basin 
and also south of the Llobregat River, including 
the southernmost part of the Vallès Basin. The 
presence of the unit is more significant in the 
southwest of the Penedès depression, where it 
occupies a central part below younger terrestri-
al and marine deposits. Towards the northeast, 
the grey marls are thinner and located along the 
southeastern margin of the basin. The unit is 
well exposed in a number of outcrops near the 
towns of Vilafranca del Penedès, Sant Sadurní 
d’Anoia, Can Rosell (Subirats), Cerdanyola or 
Rubí; some of them have yielded decapod crus-
tacean remains (Müller, 1993; Artal, 2008; Ga-
rassino et al., 2009). The Vilafranca marls con-
tain also bivalves, gastropods, echinoids, benthic 
and planktonic foraminifera, as well as remains 
of flora (Permanyer, 1982). The age of the unit is 
based on planktonic foraminifera: late Burdiga-
lian to Langhian (Macpherson, 1994). The marls 
are interpreted to have formed in an offshore 
environment (Permanyer, 1982; Cabrera et al., 
1991), although towards the top of the unit they 
must have originated in a progressively shallow-
er environment. The crab-bearing levels are lo-
cated in the middle of the sections studied and 
are attributed to the Langhian.
At the Santa Margarida site, about two kilo-
metres southwest of Vilafranca del Penedès, 
fossiliferous and intensely bioturbated yellow-
ish calcarenites, alternating with calcisiltites or 
marls, crop out. Calcisiltites and marls are more 
frequent towards the middle of the basin (to the 
northwest), whereas calcisiltites prevail towards 
the basinal margin (to the southeast). These sed-
iments are several tens of metres thick and are 
well exposed along a SW-NE strip attached to 
the Prelitoral range, south of the town of Vila-
franca, near the villages of Moja, Santa Margari-
da i els Monjos and Castellet, among others. The 
calcarenites are rich in fragments of red algae, 
planktonic and benthic foraminifera, corals, 
molluscs, echinoids, fish teeth and decapod crus-
taceans. Their age ranges from late Burdigalian 
to Langhian (Macpherson, 1994). The unit is lo-
cated in a transition zone between the carbonate 
ramp to the southeast and the open marine basin 
marls to the northwest. It is interpreted to rep-
resent distal deposits as a result of erosion and 
transport of sediments originating in the adja-
cent coralgal complex. Decapod crustacean as-
semblage at the Santa Margarida site is dominat-
ed by portunids such as Portunus monspeliensis 
(A. Milne-Edwards, 1860) and Necronectes bat-
alleri (Via, 1941). The crab-bearing calcarenites 
and marls of this unit are similar to Serraval-
lian strata in the Camp Basin, which also yield 
remains of the same crab species (Via, 1932; Ossó, 
2010). 
Repositories: MGB, Museu de Geologia de 
Barcelona-Museu de Ciències Naturals de
Barcelona (Catalonia); MV, Museu de Vila-
franca “Vinseum” (Vilafranca del Penedès, Cat-
alonia).
Systematic palaeontology
Order Decapoda Latreille, 1802
Infraorder Brachyura Latreille, 1802
Section Eubrachyura de Saint Laurent, 1980
Subsection Heterotremata Guinot, 1977
Superfamily Leucosioidea Samouelle, 1819
Family Iphiculidae Alcock, 1896
Iphiculus Adams & White, 1849
Type species - Iphiculus spongiosus Adams & 
White, 1849, by monotypy.
Iphiculus eliasi Hyžný & Gross, 2016
Figures 2A-E, 3A-E
1941 Illinae, Ebaliinae? - Vía, p. 68, pl. 10, fig. 75.
partim 1965 Petrochirus cfr. priscus Brocchi 
- Veiga Ferreira, p. 142, pl. 2, fig. 8 [non figs. 7, 9, 
11, 12 = Petrochirus priscus]
1993 Randallia? sp. - Müller, p. 12, figs. M-N.
2016 Iphiculus eliasi Hyžný & Gross, p. 266, 
figs. 2A, 3A-E, 4A-C.
104 Àlex OSSÓ, Matúš HYŽNÝ, Manu GÓMEZ, David ALBALAT & Fernando A. FERRATGES
Material and measurements (in mm): MGB 
89842 (internal mould of near-complete dorsal 
carapace): length=14.5; width=17.0; fronto-or-
bital width=6.0. MV15169 (internal mould of 
complete dorsal carapace preserving remains of 
cuticle): length=14.5; width=17.5; fronto-orbital 
width=6.0. MGB 89843 (counterpart of a male 
venter): length=8.0; width=10.5. 
Emended description: Carapace small, trans-
versely subovate in outline, widest at midlength 
(at level of posteriormost anterolateral spine), 
dorsal surface moderately convex in both di-
rections. Front not projected, bilobed, slightly 
raised, very narrow, about 0.13 of total width, me-
dially notched, strongly divergent. Orbits small, 
concave, anteriorly directed; outer orbital spine 
acute; inner orbital spine fused with frontal lobe; 
supraorbital margin with subtriangular spine, 
bounded by two open fissures. Fronto-orbital 
margin about 0.35 of total width. Lateral margins 
with 6 conical teeth; anterolateral margin with 4 
teeth, fourth being most prominent; posterolater-
al margin with 2 teeth; corners between postero-
lateral and posterior margins pointed; posterior 
margin straight, narrow, medially notched. Dor-
sal surface of carapace evenly covered with nu-
merous densely packed granules, nearly identical 
in size (when cuticular surface preserved) or with 
round concave pustules (when cuticular surface 
Fig. 2. Iphiculus eliasi Hyžný & Gross, 2016. A-D, MGB 89842 from the Langhian of Vilafranca del Penedès (Catalonia); A: 
dorsal view; B: dorsal view (digital reconstruction); C: posterior view, closeup of posterior part of carapace; D: left lateral 
view. E, MGB 89843 (male) from the Langhian of Vilafranca del Penedès (Catalonia): ventral view. Abbreviations: st = thoracic 
sternites; s = pleonal somites; t = telson. Scale bars equal 5 mm.
105On the occurrence of Iphiculus eliasi Hyžný & Gross, 2016 (Decapoda, Brachyura, Leucosioidea) from the Miocene of...
missing). Carapace surface covered evenly with 
large rounded tubercles; hepatic region with 1 
tubercle; protogastric region with one pair of 
transversely aligned tubercles in each lobe, and 
2 tubercles aligned at the basis of mesogastric re-
gion, branchial region with 3 tubercles. Carapace 
grooves absent in anterior carapace portion, well 
developed in posterior carapace portion. Gas-
tric region large, indistinctly demarcated with 
grooves. Cardiac region ovate in outline, strongly 
arched. Branchial regions broad. Intestinal re-
gion narrow. Thoracic sternum relatively wide, 
maximum width at level of fifth thoracic stern-
ite, sterno-pleonal cavity reaching end of stern-
ite 3; sternite 3 subtriangular, inverted; sternite 
4 subtrapezoidal, wider than sternite 3; sternite 
Fig. 3. Iphiculus eliasi Hyzny & Gross, 2016 in I. convexus Ihle, 1918 . A, B, D, E: Iphiculus eliasi Hyzny & Gross.: MV15169, 
from the Langhian of Santa Margarida i Els Monjos (Catalonia); A: dorsal view; B: frontal view; D: right lateral view; E: 
posterior view. C: holotype UMJGP 75.612, from the lower Badenian of Wetzelsdorf, Austria; C: dorsal view. F, G, Iphiculus 
convexus Ihle, 1918, ZRC 2009.0462 (male specimen from Vanuatu); F: dorsal view; G: ventral view. Scale bars equal 5 mm, 
except for F and G in which it is 10 mm. Photographs of F-G by P.K.L. Ng.
106 Àlex OSSÓ, Matúš HYŽNÝ, Manu GÓMEZ, David ALBALAT & Fernando A. FERRATGES
5 subrectangular transversely elongate; sternite 
6 subtrapezoidal transversely elongate; sternite 
7 subtrapezoidal, directed posteriorly, shorter 
than sternite 6. Episternite 4 laterally directed; 
episternites 5-6-7 progressively posteriorly di-
rected. Suture 3/4 laterally visible, opened; su-
tures 4/5, 5/6 and 6/7 apparently complete. Male 
pleon extremely narrow, inverted T-shaped, all 
pleonal somites free; somite 3 being widest, sub-
rectangular transversely elongate; somites 4, 5, 
and 6 subrectangular, narrowing progressively 
to the telson; telson subtriangular longitudinally 
elongate, sharp pointed, twice as long as somite 
6. Pterygostome subtrapezoidal. All ventral sur-
face, sternum, pleon, pterygostome and branchi-
ostegite densely granulate. Exognath of third 
maxilliped elongate, inner side smooth.
Remarks: Hyžný & Gross (2016) described 
a new species, Iphiculus eliasi (Fig. 3C), from 
the Middle Miocene of Austria (Steiermark). In 
their paper, Hyžný & Gross (2016, p. 268) pointed 
out that a leucosioid found in outcrops at Santa 
Margarida i Els Monjos (Alt Penedès, Catalonia), 
described and figured first as “Iliinae, Ebalii-
nae?” by Via (1941, p. 68-69, pl. 10, fig. 75) and 
subsequently as “Randallia? sp.” by Müller (1993, 
p. 12, figs. 5M-N), could be an iphiculid related 
to I. eliasi. Access to this sample of “Randallia 
sp.?” (Fig. 3A, B, D, E), housed in the Museum 
of Vilafranca (now Vinseum), has now allowed 
to conclude that, despite the different types of 
preservation as a result of different lithologies 
at outcrops, it is conspecific with the Vilafran-
ca specimens and likewise, both specimens are 
also conspecific with the Austrian one described 
by Hyžný & Gross (2016) as Iphiculus eliasi (see 
Hyžný & Gross, 2016, p. 268). Additionally, a 
small counterpart of a well-preserved male ven-
ter, recovered in the Vilafranca outcrop, is avail-
able (Fig. 2E). This exhibits the main diagnostic 
characters of the Iphiculidae, such as a very nar-
row male pleon with all somites free (Figs. 2E, 
3G; Ng et al., 2008, p. 87); this allows us to attrib-
ute it to a single iphiculid known from the area, 
I. eliasi. 
A carapace preserved in dorsal aspect from 
the Middle Miocene of Quinta da Farinheira, Lis-
boa (Portugal), described and figured by Veiga 
Ferreira (1965, p. 142-143, pl. 2, 8) as Petrochirus 
cf. priscus Brocchi, 1883, does not represent part 
of a hermit crab, but rather the carapace of a leu-
cosioid crab. Actually, the material is considered 
conspecific with Iphiculus eliasi, thus widening 
the distribution of the species further to the west.
Discussion
Müller (1993, p. 5, table 1) already point-
ed out the affinities between the Langhian and 
Serravallian decapod crustacean assemblages 
of the westernmost Proto-Mediterranean and 
roughly coeval Badenian assemblages of the 
Central Paratethys. Although he concluded that 
only 9 of 22 identified Miocene species of Cata-
lonia were reported also in the Central Parate-
thys, recent reports present further taxa that are 
present in both areas (Díaz-Medina et al., 2018; 
herein). Nevertheless, some Iberian occurrences 
are slightly younger, being of Late Miocene age 
(Díaz-Medina et al., 2017).
The presence of Iphiculus eliasi in the north-
east and southwest of Iberian Peninsula repre-
sents the westernmost (fossil) record for the ge-
nus and for the family. Extant representatives of 
the family Iphiculidae are found mainly in the 
Indo-West Pacific, in a depth range of 11 to 177 m 
(Chen, 1989; Chen & Sun, 2002), preferring mud-
dy and sandy bottoms (Galil & Ng, 2007), simi-
lar environments to that inhabited by Iphiculus 
eliasi.
Conclusions 
The ventral counterpart, preserving diag-
nostic sternal and pleonal characters, allows to 
emend the original description of Iphiculus eliasi 
and further corroborate its original systematic 
assignment. Its presence in the Middle Miocene 
of the northeastern coast of the Iberian Penin-
sula, as well as along the southwestern Iberian 
coast (Portugal), supports the circum-Mediter-
ranean distribution of decapod crustacean as-
semblages during that time interval (Gašparič & 
Ossó, 2016; Hyžný & Gross, 2016; Díaz-Medina et 
al., 2018 and references therein).
Acknowledgements
We are grateful to Quim Pastó, Mas Boquera 
(Baix Camp, Catalonia), who was among the first to 
discover the decapod crustacean assemblage at the 
Vilafranca site and brought it to our attention. Hiroaki 
Karasawa (Mizunami Fossil Museum, Gifu, Japan) 
offered the first taxonomic evaluation of the material 
as far back as 2006. Peter K.L. Ng (NUS, Singapore) 
provided photographs of extant comparative We 
thanks the staff of Vinseum (Vilafranca del Penedès, 
Catalonia) for the facilities for access to collections. 
by Alessandro Garassino (Loma Linda University, 
Loma Linda, California, USA) and Jonathan Wallaard 
(Oertijdmuseum, Boxtel, the Netherlands) greatly im-
proved the original manuscript. The research of MH 
107On the occurrence of Iphiculus eliasi Hyžný & Gross, 2016 (Decapoda, Brachyura, Leucosioidea) from the Miocene of...
was supported by VEGA 02/0136/15 and Hungarian 
Scientific Research Fund (OTKA K112708). The rese-
arch of FAF is supported by  Project E18 Aragosaurus: 
Recursos Geológicos y Paleoambientes of the gover-
nment of Aragón-FEDER.
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