GeolCarp_Vol67_No2_165

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, APRIL 2016, 67, 2, 165—176

doi: 10.1515/geoca-2016-0011

Euryhaline preferences of the decapod crab Mioplax socialis

enabled it to survive during the Badenian/Sarmatian

extinction (Miocene) in the Central Paratethys

MATÚŠ HYŽNÝ

1,2

, OLEG MANDIC

, MATHIAS HARZHAUSER

and PETER LEDVÁK

Geological-palaeontological Department, Natural History Museum Vienna, Burgring 7, 1010 Vienna, Austria; hyzny.matus@gmail.com

Department of Geology and Palaeontology, Faculty of Natural Sciences, Comenius University, Mlynská dolina, Ilkovičova 6,

842 15 Bratislava, Slovakia

The East Slovak Museum, Hrnčiarska 7, 040 01 Košice, Slovakia

(Manuscript received October 13, 2015; accepted in revised form March 10, 2016)

Abstract: Although decapod crustaceans of the Central Paratethys were diverse during the Badenian (Langhian—Early
Serravallian), a dramatic drop in their diversity occurred at the boundary with the Sarmatian. A crab Mioplax socialis is
one of the few decapods reported from the Lower Sarmatian (Mohrensternia Zone) of the Paratethys. Until now, this
species has been known from only a handful of specimens from Austria, Croatia and Bulgaria (Central Paratethys), and
its systematics and ecology remain poorly known. Here, on the basis of new specimens from the Sarmatian tuffitic clays
of the Stretava Formation (Skároš, Eastern Slovakia) we confirm that this species belongs to the subfamily
Chasmocarcininae. The diagnostic characters of the male sternum that allow this classification are reported for the first
time. The molluscan assemblage co-occurring with M. socialis demonstrate that this species tolerated conditions with
variable salinity. Its tolerance of a broad range of salinity regimes may thus explain its survival across the Badenian—
Sarmatian extinction event. Preservation of near-complete and fully articulated individuals of M. socialis suggests calm
conditions and short residence times on the sediment-water interface.

Key words: Sarmatian, Crustacea, Mollusca, stratigraphy, euryhaline assemblage.

Introduction

Decapod  crustaceans  are  important  components  of  marine
benthic associations of the continental shelf and slope (Bou-
dreau & Worm 2012; Lee 2015). Their evolutionary success
is  connected  with  several  major  diversification  events;  one
of  them  occurred  during  the  Miocene  (Schweitzer  2001;
Feldmann & Schweitzer 2006). This Miocene diversification
was  enhanced  by  the  biogeographical  differentiation  of  the
Western Tethys into two different palaeogeographical areas,
including  the  circum-Mediterranean  and  the  Paratethys
(Rögl  1998,  1999;  Popov  et  al.  2004;  Harzhauser  &  Piller
2007). Numerous decapod species have been recorded espe-
cially  in  the  Central  Paratethys,  many  of  them  being  Para-
tethyan endemites.

One of the latest marine decapod associations of the Cen-

tral Paratethys occurs in the lowermost Sarmatian strata (up-
per  Serravallian)  at  Devínska  Kobyla  in  the  Slovak  part  of
the Vienna Basin (Hyžný 2012; Hyžný & Hudáčková 2012;
Hyžný  et  al.  2012).  The  stenohaline  communities  seem  to
disappear  at  the  Badenian/Sarmatian  boundary  (Harzhauser
& Piller 2007; Studencka & Jasionowski 2011) or during the
earliest Sarmatian (Hyžný et al. 2012), depending on the re-
gional  differences  in  environmental  conditions.  They  were
replaced  by  homogeneous  euryhaline  biota,  most  of  which
were  endemic  to  the  region  (Rögl  1998,  1999;  Popov  et  al.
2004).  However,  occurrences  of  a single  crab  species,
Mioplax socialis Bittner, 1884, were reported from the Lower

Sarmatian of Waldhof, Austria (Glaessner 1928), Kochava,
Bulgaria (Müller 1979) and Skároš, Slovakia (Hyžný & Led-
vák 2014); all these occurrences postdate the Badenian deca-
pod associations from Devínska Kobyla (Hyžný et al. 2012).
Mioplax socialis, until now known only from the Paratethys,
seems to be such a Lower Sarmatian endemic species adapted
to euryhaline conditions. This species probably represents
the last marine crab of the Central Paratethys.

The type collection of M. socialis has not been studied

since its description in the 19

century and only a few speci-

mens of this species were known. As a consequence, all taxo-
nomic  treatments  have  been  largely  based  on  figures  and
original  description  of  Bittner  (1884);  later  reports  by
Glaessner (1928) and Müller (1979) did not add much to the
knowledge  of  this  crab.  Recently,  new  Mioplax  specimens
were found in the Eastern Slovakia (Hyžný & Ledvák 2014).
In this study, we analyse the co-occurring  molluscan-bryo-
zoan assemblage, revisit and re-figure the type material, and
document  the  environmental  preferences  of  this  last  marine
crab of the Central Paratethys.

Geological setting

The macrobenthic assemblage documented here is derived

from a locality situated on the western slope of the Slanské
Mts., approximately 1 km SE from the Skároš village (GPS
N 48°34 49.58”, E 21°24 19.43”), Eastern Slovakia

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(Fig. 1b).  In  the  right  bank  of  the  creek,  a  sequence  of  cal-
careous  tuffitic  clays  of  the  Stretava  Formation  (Vass  &
Čverčko  1985)  is  exposed,  locally  with  thin  layers  of  pu-
mice.  The  studied  section  is  approximately  2.5  m-thick.  At
the  base  and  on  the  top  of  the  section,  tuffs  are  exposed
(Fig. 1c). They are largely composed of well-sorted volcano-
clastics  and  pumice.  The  tuffs  do  not  contain  any  fossil
macrofauna.  Among  the  tuffs,  several  layers  of  calcareous
tuffitic clay with a total thickness of approximately 1.5 m are
exposed. Calcareous clays (Fig. 1e) contain molluscs, crabs
(a  monospecific  suite  of  Mioplax  socialis)  and  bryozoans.
On the basis of the molluscan assemblage, the exposed strata
at the locality are estimated to be of Lower Sarmatian of age
(Mohrensternia Zone).

Material and methods

The specimens of Mioplax socialis described by Bittner

(1884)  and  Glaessner  (1928)  and  newly  collected  material
from  several  localities  in  Eastern  Slovakia,  namely  Skároš,
Trstené  pri  Hornáde  and  Nižná  Myš a.  More  extensive
sampling  at  one  of  these  localities,  Skároš  (see  above;
Fig. 1), also provided a molluscan assemblage that allows us

to evaluate the palaeoenvironmental conditions (see Discus-
sion).

Both, molluscs and decapods, were preserved largely as

internal moulds. The material (1733 g) comprised presorted
sediment samples with remains of macrofauna, including
83 specimens of molluscs (bivalves and gastropods) and
10 specimens of brachyurous crabs.

The aragonitic mollusc shells were completely leached;

therefore, casts have been produced using two component
vinyl polysiloxane (VPS), allowing an insight into original
morphology. The specimens of decapods were photographed
dry and uncoated or coated with ammonium chloride; for de-
tails see figure captions.

The repositories of specimens illustrated or referred to be-

low are as follows:
GBA – Geological Survey, Vienna (Austria)
KGP-MH  –  “Matúš  Hyžný  collection”  at  Department  of
Geology  and  Palaeontology,  Faculty  of  Natural  Sciences,
Comenius University, Bratislava (Slovakia)
NHMW  –  Geological-palaeontological  Department,  Natu-
ral History Museum, Vienna
UMJGPA – Geological-palaeontological Department, Uni-
versalmuseum Joanneum, Graz (Austria)
VSM – The East Slovak Museum, Košice (Slovakia)

Fig. 1. Geographical position (a) and geological setting (b) of the surroundings of the locality Skároš. Legend: A – andesites and dacites
(Sarmatian—?Pannonian); B – andesitic pyroclastics and epiclastics (Sarmatian—?Pannonian); C – rhyolitic tuffs and tuffites (Lower Sar-
matian); D – clays and tuffites of the Stretava Formation (Lower and Middle Sarmatian); E – Quaternary deposits. c – Outcrop in the
Stretava Formation. Fossiliferous tuffitic clays (demarcated by white lines) are exposed between volcanoclastics above and below them.
d—e – Detailed view on the clays with Lower Sarmatian macrofauna. Map (b) modified after Kaličiak et al. (1996). Photographs in c and e
by MH, d by PL.

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Systematic Paleontology

Identifications of molluscs are based on Kolesnikov

(1935),  Papp  (1954,  1974),  Boda  (1959),  Kojumdgieva
(1969),  Švagrovský  (1971),  Nevesskaya  et  al.  (1993),
Schultz (2001, 2003, 2005), Harzhauser & Kowalke (2002),
Kowalke & Harzhauser (2004), and Lukeneder et al. (2011).
Mollusc  synonymy  lists  include  only  the  first  descriptions
and the most recent or important references. Works of Ng et
al.  (2008),  De  Grave  et  al.  (2009)  and  Schweitzer  et  al.
(2010)  are  followed  for  higher  classification  of  Mioplax
socialis crabs. Synonymy of M. socialis is extensive.

Phylum Mollusca Linnaeus, 1758

Class Bivalvia Linnaeus, 1758

Subclass Pteriomorphia Beurlen, 1944

Superfamily Mytiloidea Rafinesque, 1815

Family Mytilidae Rafinesque, 1815

Genus Musculus Röding, 1798

Type species: Mytilus discors Linnaeus, 1767; subsequent

designation by Iredale (1915); Recent, European Seas.

Musculus

sarmaticus (Gatujev, 1916)

Fig. 2b

*1916 Modiolus (Brachydontes) sarmaticus m. — Gatujev, p. 148—149,

pl. 12, fig. 1.

2001 Musculus (Musculus) sarmaticus (Gatujev, 1916) — Schultz,

p. 109—114, pl. 8, fig. 16.

Material examined: Five small sized (maximum length:

11.4 mm), disarticulated specimens preserved as interior and
exterior valve imprints.

Remarks: They are of subtrigonal outline, flattened, with

weak radial riblets on central and dorsal shell exterior, interior
margin shows minute crenulations.

Occurrence: Sarmatian of the Central Paratethys; Konkian

to Bessarabian of the Eastern Paratethys (Schultz 2001).

Subclass Heterodonta Neumayr, 1884

Superfamily Cardioidea Lamarck, 1809

Family Cardiidae Lamarck, 1809

Subfamily Lymnocardiinae Stoliczka, 1870

Genus Obsoletiforma Kolesnikov, 1948

Type species: Cardium vindobonense Laskarev, 1903, origi-
nal designation. Miocene, Europe.

Obsoletiforma

cf. vindobonensis (Laskarev, 1903)

Figs. 2c—e

* 1903 Cardium vindobonense Partsch — Laskarev, p. 79, 141, pl. 3,

figs. 14—19.

2003 Obsoletiforma (Obsoletiforma) vindobonensis vindobonensis

(Laskarew, 1903) — Schultz, p. 585—595, pl. 87, figs. 1—2.

Material: 16 disarticulated and 7 articulated specimens

are available, with a maximum length of 11.2 mm.

Remarks: Outline is anteriorly rounded, posteriorly trun-

cated, transversal keel is moderately prominent. Ribs (~16

on the anterior to central and 8 on posterior shell portion) are
flattened and smooth except for delicate scales present on
anterior rib tops.

Shells are identical in size and in sculpture with specimens

from  the  Mohrensternia  Zone  of  Aspersdorf/Hollabrunn  in
Lower Austria (Papp 1954, 1974; Mandic et al. 2008). Both,
however,  lack  the  dense  transversal  scale  ornamentation  on
rib tops typical for Obsoletiforma vindobonensis (Laskarev,
1903), but absent in O. lithopodolica (du Bois 1831), which
seems  to  have  in  contrast  a  different  outline  (Kojumdgieva
1969; Nevesskaya et al. 1993). Both latter species were ori-
ginally described from the Carpathian Foredeep, demanding
a  thorough  revision  of  their  true  relationship  to  the  Panno-
nian  Basin  Obsoletiforma.  Yet,  the  preservation  of  the  stu-
died  material  does  not  allow  more  precise  species-level
identification at the moment.

Occurrence: Badenian to Sarmatian of the Central Para-

tethys, Konkian to Lower Bessarabian of the Eastern Para-
tethys (Schultz 2003).

Superfamily Tellinoidea Blainville, 1814

Family Semelidae Stoliczka, 1870

Genus Abra Lamarck, 1818

Type species: Mactra tenuis Montagu, 1803; by subse-

quent designation (Herrmannsen 1846); Recent, NE Atlantic
and Mediterranean Sea.

Abra reflexa

(Eichwald, 1830)

Fig. 2a

*1830 Donax reflexa, m. — Eichwald, p. 208.
2005 Abra (Syndosmya) reflexa (Eichwald, 1830) — Schultz, p. 746—748,

pl. 102, figs. 1—2.

Material: 3 articulated and 17 disarticulated specimens.
Remarks: Flattened, smooth shell, subtrigonal in outline

rounded anteriorly, and pointed posteriorly. With maximum
shell length of 18.4 mm this is the largest mollusk species
present in the samples.

Occurrence: Badenian to Lower Sarmatian of the Central

Paratethys, Konkian to Bessarabian of the Eastern Paratethys
(Kojumdgieva 1969; Papp et al. 1974; Studencka et al. 1998;
Schultz 2003).

Class Gastropoda Cuvier, 1795

Subclass Vetigastropoda Salvini-Plawen, 1980

Superfamily Trochoidea Rafinesque, 1815

Family Trochidae Rafinesque, 1815

Genus Gibbula Risso, 1826

Type species: Trochus magus Linnaeus, 1758, by subse-

quent designation (Herrmannsen 1847). Recent, Mediterra-
nean Sea.

Gibbula

cf. guttenbergi (Hilber, 1897)

Fig. 2h

*1897 Trochus Guttenbergi Hilber, species nova — Hilber, p. 194—195,

figs. 7—8.

1974 Calliostoma (F[eneoniana].) guttenbergi (Hilber) — Papp, p. 325,

pl. 1, figs. 1—3.

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Material: With 17 specimens it is relatively abundant in

the studied material.

Remarks: With only 6.5 mm maximum shell height this is

a very small trochoid species. The trochiform shell comprises
about 6 slightly convex whorls continuously increasing in
size, ornamented by numerous delicate spiral ribs. Although
highly similar to specimens from the Mohrensternia-beds of
Waldhof in the Styrian Basin (e.g. Papp 1954), representing
its type locality, complete specimens are missing and there-
fore the identification remains uncertain. As shown by Harz-
hauser & Kowalke (2002), the Sarmatian species formerly
treated as Calliostoma Swainson, 1840 have to be placed in
Gibbula Risso, 1826 based on protoconch morphology.

Occurrence: Lower Sarmatian of the Central Paratethys

(Papp et al. 1974).

Gibbula

cf. blainvillei (d’Orbigny, 1844)

Fig. 2f

*1844 Trochus Blainvillei d’Orb., 1844— d’Orbigny, p.446—447, pl. 2,

figs. 3—5.

1969 Gibbula (Gibbula) blainvillei (d’Orbigny, 1844) — Kojumdgieva,

p. 68, pl. 24, fig. 20, pl. 25, fig. 1.

Material: A single depressed conical shell of 16.8 mm

diameter.

Remarks: A shell with wide, moderately convex last

whorl, flaring keel and smooth shell surface except for a sin-
gle spiral cord is available. It seems to represent this general-
ly rare species. Gibbula papilla (Eichwald, 1853), which has
a comparable shape, is smaller. Both species are known so
far only from Bessarabian deposits. This species group, how-
ever, has its roots in the Early Sarmatian Gibbula sopronen-
sis (Papp, 1954), which is higher and has flat whorls. Due to
this stratigraphic gap and the poor preservation we prefer to
identify the specimen in open nomenclature.

Occurrence: Bessarabian of the Carpathian Foredeep and

the Black Sea Basin (Kojumdgieva 1969); this is its first
record from the Sarmatian of the Pannonian Basin System,
namely the Central Paratethys.

Subclass Caenogastropoda Cox, 1960

Superfamily Cerithioidea Fleming, 1822

Family Batillariidae Thiele, 1929

Genus Granulolabium Cossmann, 1889

Type species: Cerithium plicatum Bruguière, 1792, by

original designation. Early Miocene, France.

Granulolabium bicinctum

(Brocchi, 1814)

Fig. 2g

*1814 Murex bicinctus nob.— Brocchi, p. 446, pl. 9, fig. 13.
2013 Granulolabium bicinctum (Brocchi, 1814) — Landau et al., p. 45,

pl. 4, figs. 3—4. [cum. syn.]

Material: Only four specimens are present with maximum

height of 18.0 mm.

Remarks: Shell comprises about 9 whorls, ultimate whorl

attains 42 % of shell height. Sculpture, fully developed in last
three whorls only, comprising three spiral keels and about

10 axial folds producing nodes at intersection with the ribs.
Shell  base  bears  two  strong  spiral  keels.  The  species  origi-
nated in the Early Miocene of the Eastern Atlantic, Mediter-
ranean  and  Paratethys  domain  (Landau  et  al.  2013),  and
represents  a  survivor  of  the  Badenian/Sarmatian  extinction
event.  G.  pictum  (Defrance  in  Basterot,  1825),  a  name  pre-
viously  widely  used  for  the  Sarmatian  records,  is  its  junior
synonym (Landau et al. 2013).

Occurrence: Early Miocene of NE Atlantic Ocean, Early

to Middle Miocene of Paratethys, and Early Miocene to
Pliocene of (Proto-)Mediterranean Sea (Landau et al. 2013).

Superfamily Rissooidea Gray, 1847

Family Rissoidae Gray, 1847

Subfamily Mohrensterniinae Korobkov, 1955

Genus Mohrensternia Stoliczka, 1868

Type species: Rissoa angulata Eichwald, 1853, subse-

quent designation (Nevill, 1885). Badenian and Early Sar-
matian of Central and Eastern Europe.

Remarks: Mohrensternia is quite frequent in the samples

(13 specimens) but a species level identification is difficult
due to the poor preservation. Beside Mohrensternia pseudo-
inflata Hilber, 1897 the samples also seem to contain
Mohrensternia inflata (Hörnes, 1856).

Mohrensternia

cf. pseudoinflata Hilber, 1897

Fig. 2i—j

*1897 [Mohrensternia] pseudinflata Hilb. — Hilber, p. 201.
1971 Mohrensternia pseudinflata Hilber, 1897 — Švagrovský, p. 285—

287, pl. 44, figs. 6—9.

Material: At least one specimen.
Remarks: Shell minute, with maximum height of 8 mm,

ovate  conic,  with  6  moderately  inflated,  slightly  angular
whorls;  last  whorl  attaining  68%  of  shell  height.  Sculpture
consists  of  prominent  axial  ribs  with  delicate  spiral  threads
in the interspaces (Fig. 2i). This species differs from M. in-
flata in the presence of spiral threads and in the more conical
outline  and  the  distinctly  less  inflated  and  angulated  last
whorl.  Mohrensternia  pseudosarmatica  Friedberg,  1923
(sensu  Švagrovský  1971)  is  similar  in  sculpture  but  has
deeper sutures and strongly convex whorls.

Occurrence: Lower Sarmatian of the Central and Eastern

Paratethys (Švagrovský 1971; Papp et al. 1974). Kowalke &
Harzhauser (2004) reported it from the Konkian, which is the
Eastern Paratethys equivalent of Late Badenian.

Phylum Arthropoda von Siebold, 1848

Class Malacostraca Latreille, 1802

Order Decapoda Latreille, 1802

Infraorder Brachyura Latreille, 1802

Section Eubrachyura de Saint Laurent, 1980

Subsection Heterotremata Guinot, 1977

Superfamily Goneplacoidea MacLeay, 1838

Family Chasmocarcinidae Serène, 1964a

Subfamily Chasmocarcininae Serène, 1964a

Genus Mioplax Bittner, 1884

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Type species: Mioplax socialis Bittner, 1884, by monoty-

py.

Diagnosis: As for the type species.
Remarks: The genus was erected to accommodate a sin-

gle  species,  Mioplax  socialis,  from  the  Badenian  strata  of
present day Croatia (Bittner 1884). Later, it has also been re-
ported  from  the  Sarmatian  of  Austria  (Glaessner  1928)  and
the  genus  was  placed  within  the  subfamily  Goneplacinae
MacLeay,  1838,  the  family  Goneplacidae  MacLeay,  1838
(Glaessner  1969).  Karasawa  &  Kato  (2003)  argued  that  the
presence of a small orbit and long, slender deflexed fingers
of chelipeds strongly suggest assignment to the Chasmocar-
cininae Serène, 1964a. The subfamily was elevated to family
level  by  Karasawa  &  Schweitzer  (2006),  and  this  was  also
followed by subsequent workers (see classifications by Ng et
al. 2008; De Grave et al. 2009; Schweitzer et al. 2010). Pre-
sence of the supplementary coxosternal plate on male sternite 8
(Guinot  et  al.  2013),  documented  herein  for  the  first  time
(Fig. 4f), and possession of male pleonal somites 3—5 fused
(Feldmann et al. 2010) further strengthen inclusion of the ge-
nus  into the Chasmocarcinidae.  Mioplax is similar to other
chasmocarcinids, however, it possesses a well-developed an-
terolateral spine, which is not present in any other genus of
the family. The genus is rather close to Collinsius Karasawa,
1993,  Falconoplax  Van  Straelen,  1933,  Orthakrolophos
Schweitzer  &  Feldmann,  2001,  and  Styrioplax  Glaessner,

1969 (all four genera are known exclusively as fossils), but
can  be  immediately  distinguished  from  them  based  on  the
presence of the aforementioned anterolateral spine. Another
exclusively  fossil  genus,  Gillcarcinus  Collins  &  Morris,
1978, is characterized by mesobranchial ridges, quite untypi-
cal  for  any  other  chasmocarcinid.  Similarly,  representatives
of  extant  genera,  Camatopsis  Alcock  &  Anderson,  1899,
Chasmocarcinus  Rathbun,  1898,  Chasmocarcinops  Alcock,
1900,  and  Hephthopelta  Alcock  1899,  do  not  have  distinct
anterolateral spine (e.g., Rathbun 1898; Serène 1964b).

Mioplax socialis

Bittner, 1884

Figs. 3—4

1884 Mioplax socialis Bittner, p. 23, pl. 2, figs. 3a—f.
1897 “Krabbe, Fam. Catometopa”. — Hilber, p. 109.
1928 Mioplax socialis Bittner. — Glaessner, p.194.
1929 Mioplax socialis Bittner. — Glaessner, p. 258.
1929 Mioplax socialis Bittner. — Lőrenthey & Beurlen, p. 258, pl. 16,

fig. 10.

?1940 Potamon hungaricum Körössy, fig. 9.
?1955 Pseudopotamon hungaricum (Körössy) — Bott, p. 310.
?1969 Pseudopotamon hungaricum (Körössy) — Bott, p. 272.
1969 Mioplax socialis Bittner. — Glaessner, p. R524, fig. 333.1.
1979 Mioplax cf. socialis Bittner. — Müller, p. 6, pl. 3, fig. 3.
2003 Mioplax socialis Bittner. — Karasawa & Kato, table 8.
2010 Mioplax socialis Bittner. — Schweitzer et al., p. 133.
?2010 Potamon? hungaricum Körössy. — Klaus & Gross, p. 49, fig. 5D.
2014 Mioplax socialis Bittner. — Hyžný & Ledvák, p. 28, figs. 1—2.

Fig. 2. Molluscs identified in the studied samples from the Stretava Formation exposed at Skároš, Slovakia. a – Abra reflexa (Eichwald,
1830), left valve exterior (NHMW 2015/0400/0001); b – Musculus sarmaticus (Gatujev, 1916), right valve exterior (NHMW 2015/0400/
0002); c—e – Obsoletiforma cf. vindobonensis (Laskarev, 1903), (c) right valve exterior (NHMW 2015/0400/0003), d – articulated shell
dorsal view (NHMW 2015/0400/0004), e – left valve exterior (NHMW 2015/0400/0005); f – Gibbula cf. blainvillei (d’Orbigny, 1844)
(NHMW 2015/0400/0006); g – Granulolabium bicinctum (Brocchi, 1814) (NHMW 2015/0400/0007); h – Gibbula cf. guttenbergi
(Hilber, 1897) (NHMW 2015/0400/0008); i—j – Mohrensternia cf. pseudoinflata Hilber, 1897 (NHMW 2015/0400/0009). All photo-
graphs by OM.

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Emended diagnosis: Carapace flat, square, only cardiac

region marked; front wide, straight, single well-developed
anterolateral spine directed forward.

Type material (chosen herein from the syntype collec-

tion): Radoboj, Croatia: lectotype GBA 2009/0014/0029/01
(a  near-complete  specimen  with  preserved  carapace,  both
chelipeds  and  partly-preserved  walking  legs),  paralectotype
GBA/0014/0029/02  (a  female  specimen  in  ventral  aspect
with partially preserved chelipeds and  left third maxilliped),
paralectotype  GBA/0014/0029/03  (a  specimen  in  dorsal  as-
pect with preserved carapace and left cheliped), and paralec-
totype  GBA/0014/0029/04  (a  male  specimen  in  ventral
aspect with partially preserved pereiopods).

Additional material: Waldhof, Austria: UMJGPA 4695

(carapace with partially preserved venter); Skároš, Slovakia:
NHMW  2015/0401/0001—0002  (two  near-complete  indi-
viduals in dorsal aspect), NHMW 2015/0401/0003 (male in-
dividual  in  ventral  aspect),  NHMW  2015/0404/0004—0005,
KGP-MH  SK-001—008  and  VSM/P-1131;  Trstené  pri
Hornáde,  Slovakia:  VSM/P-1132;  Nižná  Myš a,  Slovakia:
VSM/P-1133.

Description: Carapace sub-rectangular, slightly wider

than  long,  widest  at  midlength;  carapace  transversely  and
longitudinally  convex,  with  steep  lateral  margins.  Front
straight, with faint median notch; orbits shallow, orbital mar-
gins  entire;  anterolateral  margin  with  one  prominent  spine
directed forward and separated with a notch towards the or-
bit;  posterolateral  margin  with  concave  re-entrants  for
pereiopods  5;  posterior  margin  straight,  rimmed.  Carapace
surface  mostly  smooth,  tiny  tubercles  present  at  carapace

margins; most regions poorly defined; gastric region well-
defined, posterior border delimited by cervical groove; cardiac
region well-defined, sub-trapezoid in outline, slightly raised
centrally, posterior border poorly defined.

Buccal frame narrowing anteriorly; third maxillipeds not

closing  buccal  cavity;  merus  of  third  maxilliped  endopod
suboval, about half the length of ischium, palp (carpus-pro-
podus-dactylus)  distal;  third  maxilliped  exopod  not  pre-
served. Sternum wide, all sutures interrupted; sterno-pleonal
cavity  reaching  anterior  of  sternite  4;  sternite  8  visible  in
ventral view; overlying posterior part of sternite 7. Sternite 8
in males with a groove in the middle covered by intercalated
supplementary coxosternal plate, coxosternal plate as long as
sternite 8. Male abdomen not entirely filling space between
coxae of pereiopods 5; male pleonal somites 3—5 fused, nar-
rowing slightly posteriorly, telson triangular.

Chelipeds (pereiopods 1) unequal, right chela stout and

larger than left chela; carpus quadrate, propodus sub-quad-
rate with smooth surface, fingers long and slender. Pereio-
pods 2—5 achelate, long and slender.

Remarks: The material from Skároš fully conforms to the

type material from Radoboj, Croatia (Bittner 1884). The dif-
ference is only in the type of preservation. Whereas the type
series  is  represented  by  three-dimensional  preservation  of
crabs within the concretions (Fig. 3a—b), specimens from the
tuffitic  clays  of  the  Stretava  Formation  are  flattened
(Fig. 3d—h)  and  in  many  aspects  they  resemble  another
Miocene  chasmocarcinid  from  the  Paratethys,  Styrioplax
exiguus  (Glaessner,  1928)  reported  from  the  Karpatian
of Cerová-Lieskové,  Slovakia  (Hyžný  &  Schlögl  2011,

Fig. 4. Mioplax socialis Bittner, 1884. a—c – Paralectotype from Radoboj, Croatia (GBA 2009/0014/0029/02). Female sternum (a—b) with
preserved third maxilliped (c). d—f – Male individual in ventral aspect with complete sternum and pleon (d) from Skároš, Slovakia
(NHMW 2015/0401/0003). Note the presence of coxosternal plate (f). Numbers in b and e denote sternites. The specimens in a—c were
coated with ammonium chloride prior to photography. The scale bar equals 5 mm (all except f) and 1 mm (f). All photographs by MH.

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text-fig. 13). Similarly, specimens of Mioplax socialis
reported by Glaessner (1928) from Waldhof (Austria) exhibit
a flattened carapace (Fig. 3c).

All the specimens possess one anterolateral spine. Its pre-

sence is recognizable due to the indent (or notch) towards the
front. The spine itself is not always preserved in its entirety,
frequently it is broken (e.g. Fig. 3a, b, d). However, its base
is  invariably  present  in  all  specimens.  The  well-preserved
spine  is  clearly  visible  in  the  Waldhof  specimen  (Fig.  3c).
The  specimen  from  Kochava  (Bulgaria)  is  crushed  and
shows a poorly preserved anterolateral margin (Müller 1979,
pl. 3, fig. 3), therefore, the diagnostic spine is not preserved.
The morphology of the carapace, however, clearly indicates
that this specimen belongs to the subfamily Chasmocarcininae
and  given  its  Sarmatian  age  there  is  little  doubt  about  its
assignment to Mioplax socialis.

The male pleon is described and figured here for the first

time  for M.  socialis  (Fig.  4d—e).  Bittner  (1884)  reported  on
only one poorly preserved male specimen, all others were fe-
males.  The  male  pleon  of  M.  socialis  is  distinctly  broader
than  that  in  Styrioplax  exiguus  (Glaessner,  1928)  (see  Gaš-
parič & Hyžný 2015, figs. 10D—E, 11B), another chasmocar-
cinid crab from the Paratethys. This species, however, occurs
in  the  Karpatian  and  Lower  Badenian  strata  (Hyžný  &
Schlögl 2011; Gašparič & Hyžný 2015).

Apart from the material from Waldhof, Glaessner (1928)

also reported two dactyli from the Sarmatian of Hernals
(Vienna Basin) which he questionably attributed to M. socialis.

Based on two specimens from the Lower Sarmatian deposits

of  the  Stretava  Formation  exposed  at  Ždaňa  (Slovakia)
Körössy  (1940)  described  two  crab  specimens  as  Potamon
hungaricum. Klaus & Gross (2010) in their synopsis of Eu-
ropean  fossil  freshwater  crabs  provided  a  brief  overview  of
this taxon and re-printed an original (rather poor) photograph
of the holotype. They considered the species unlikely to rep-
resent a potamid crab. Not only the long and narrow legs ar-
gue  against  the  assignment  to  the  genus  Potamon  Savigny,
1816, but the material comes from marine to brackish pelites
of the Stretava Formatian which are unlikely to contain arti-
culated  remains  of  freshwater  crabs.  The  type  material  of
P. hungaricum is lost (Müller, pers. comm. in Klaus & Gross
2010) and the comparison with Mioplax socialis is thus not
possible. However, according to Körössy (1940), the front of
P.  hungaricum  is  straight  and  typical  also  of  Mioplax.  The
relatively small carapaces of P. hungaricum (max. carapace
width  14  mm)  also  fit  within  the  size-range  of  M.  socialis.
Both  sedimentological  settings  (pelites  of  the  Stretava  For-
mation) and stratigraphy (Early Sarmatian), are shared by the
occurrences of P. hugaricum from Ždaňa and M. socialis of
Skároš.  Moreover,  both  localities  are  only  few  kilometres
apart.  Finally,  Zlinská  &  Fordinál  (1995)  mentioned  the
presence of crab claws that may belong to M. socialis from
the same formation at Slanská Huta.

Occurrence: Mioplax socialis has been reported from the

Badenian  of  Radoboj  in  Croatia  (Bittner  1884),  and  Lower
Sarmatian  of  Waldhof  (Mohrensternia  beds)  in  Austria
(Glaessner  1928),  Kochava  in  Bulgaria  (Müller  1979),  and
Skároš, in eastern Slovakia (Hyžný & Ledvák 2014; herein).
Müller (1979, p. 6) also mentioned occurrences in Hungary

(boreholes in the vicinity of Nagybózsva and Mány), how-
ever, he did not supply any figure. Occurrences of Potamon
hungaricum from the Lower Sarmatian of Ždaňa and Slanská
Huta in eastern Slovakia seem to belong to M. socialis.

Discussion

Stratigraphy: A poorly diverse, euryhaline-type mollus-

can-assemblage  from  Skároš,  dominated  by  cardiids,  se-
melids,  rissoids,  and  gibbulids,  implies  its  Sarmatian
(Middle  Miocene)  age  (Papp  et  al.  1974).  In  addition,  the
presence  of  Mohrensternia,  allows  a  clear  correlation  with
the  Lower  Sarmatian  Mohrensternia  Zone  (Papp  1954;
Kowalke  &  Harzhauser  2004).  This  zone  represents  the
lowermost  interval  of  the  Sarmatian  ecostratigraphic  zona-
tion in the Central Paratethys and correlates with the time in-
terval of 12.7 to 12.4 Ma (Harzhauser & Piller 2004).

At the genus level, all identified molluscan taxa represent

survivors  of  the  Badenian-Sarmatian  extinction  event
(Harzhauser & Piller 2007; Studencka & Jasionowski 2011).
At  the  species  level  and  in  the  Central  Paratethys  domain,
Obsoletiforma  vindobonensis  is  restricted  to  the  Sarmatian,
Gibbula guttenbergi and Mohrensternia pseudoinflata to the
Early Sarmatian, whereas the other three species (Musculus
sarmaticus,  Abra  reflexa,  and  Granulolabium  bicinctum)
existed already during the Badenian (Papp 1974; Studencka
et al. 1998; Landau et al. 2013). Similarly, the crab compo-
nent of the association, Mioplax socialis, is known not only
from the Early Sarmatian (Glaessner 1928; Müller 1979), but
also from the Badenian strata as documented by occurrence
of marine fauna with Anadara diluvii (Lamarck), Nucula cf.
mayeri  (Hörnes),  Perna  aquitanica  (Mayer),  Psammotreta
aff. papyracea (Chemnitz), and  Ptychidia cf. vindobonensis
(Handmann) at the type locality Radoboj, NW Croatia (Bitt-
ner 1884). Moreover, Bittner (1884) expressively stated that
the  crab-bearing  strata  probably  underlie  the  Leitha  lime-
stone (Badenian in age). Glaessner (1928, p. 194) expressed
some doubts about the age of the specimens from Radoboj.
The euryhaline preferences of Mioplax socialis would explain
why the crab has been found both in the Badenian and Sarma-
tian strata without questioning the statement of Bittner (1884).

Environment and ecology: Taxonomic abundances at

Skároš  are  rather  evenly  distributed,  with  a  mixture  of  sus-
pension-feeders and deposit-feeders, and show that the com-
munity  was  heavily  dominated  by  a  single  species  (Fig.  5).
Such  evenness  may  imply  stable  environmental  conditions,
without  strong  fluctuation  of  chemical  or  physical  parame-
ters such as salinity, oxygen, or water energy, although mul-
tiple  other  ecologic  and  taphonomic  factors,  other  than
environmental stability affect the community structure (Kid-
well & Bosence 1991). The abundant rissoids point to calm
littoral conditions (Kowalke & Harzhauser 2004). The thin-
shelled generalistic species Abra reflexa occurs in estuarine
settings  during  the  Sarmatian  (Mandic  et  al.  2008),  but
monospecific occurrences are reported only from calm, oxy-
gen  poor,  and  deeper  water  settings  (Studencka  &  Jasio-
nowski  2011).  In  contrast,  Granulolabium  bicinctum  is
typically dominant on algal mats in lower intertidal settings

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(Harzhauser & Kowalke 2002; Lukeneder et al. 2011; Landau
et al. 2013). We suggest the assemblage from Skároš is most
likely derived from the subtidal zone, close to the shore, but
at least beneath the fair-weather wave base.

With the exception of complete aragonite leaching and mi-

cro-fractures (originating from post-depositional compaction
processes), the shells are relatively well-preserved, the frag-
mented shells are rare. The proportion of articulated bivalve
shells is moderate to low (44 % of cardiids, 18 % of semelids,
and  no  mytilids  are  articulated).  Among  the  articulated
shells, the fully open individuals in butterfly position domi-
nate, whereas closed individuals are rare. In particular, only
one  such  cardiid  shell  is  present  in  the  samples,  suggesting
that shells are basically never found in their living position.
In contrast, four cardiid and one semelid shells show butter-
fly  position.  Shells  are  oriented  concordantly  with  the  bed-
ding  planes.  Such  preservation  supports  a  previous
interpretation of a calm depositional setting and low degree
of  benthic  mixing  due  to  bioturbation  (Kidwell  &  Bosence
1991).  Thus,  after  their  natural  death,  the  shells  were  gra-
dually covered by pelitic sediment. As suggested by the fre-
quent  articulated  shells,  this  preservation  occurred  without

previous dragging from the sea bottom by currents or wave
action.  Mioplax  socialis  crabs  are  commonly  preserved  as
fully articulated individuals which corroborates the interpre-
tation  of  low-energy  conditions  with  a  relatively  short  resi-
dence  time  on  the  sediment-water  interface.  Bodies  of
decapod  crustaceans  decompose  relatively  quickly  (Schäfer
1951;  Plotnick  1986;  Plotnick  et  al.  1988;  Stempien  2005;
Mutel et al. 2008); a calm depositional setting coupled with
burial without subsequent physical disturbance is a prerequi-
site for preservation of complete or near-complete specimens
(Müller et al. 2000, fig. 23).

This interpretation of rather stable conditions some dis-

tance  from  the  high-energy  environments  is  in  accordance
with  the  monospecific  presence  of  the  bryozoans
Schizoporella  cf.  tetragona  (Reuss,  1847),  which  is  com-
mon  in  the  investigated  samples  (see  also  Filipescu  et  al.
2014; Fig. 6).

Conclusions

With the exception of some lowermost Sarmatian deca-

pods occurring in the Vienna Basin, Mioplax is the sole de-
capod crab occurring in the Lower Sarmatian of the Central
Paratethys  (Mohrensternia  Zone).  The  molluscan  assem-
blage  from  the  tuffitic  clays  of  the  Stretava  Formation  ex-
posed in Eastern Slovakia (Central Paratethys) demonstrates
that  M.  socialis  was  a  euryhaline  species,  which  was  likely
the  reason  why  it  survived  the  Badenian/Sarmatian  extinc-
tion event (Harzhauser & Piller 2007). Preservation of near-
complete and articulated individuals at virtually all localities
with  M.  socialis  suggests  that  the  species  inhabited  a  calm,
low-energy setting and most probably environmental condi-
tions without a strong fluctuation in oxygen or water energy.

Acknowledgements:  We  are  thankful  to  Štefan  Miňo  for
help during fieldwork. Róbert Zajac provided information of
crab-bearing layers during the initial field studies and photo-
graphed  specimens  deposited  in  East  Slovak  Museum  (Ko-
šice). For sharing his bryozoan expertise, we are indebted to
Björn  Berning  (OÖ  Landesmuseum  Linz).  The  reviewers
(Rok Gašparič and Hiroaki Karasawa) and an associate edi-

Fig. 5. Chart showing the overall abundance of molluscs in studied
samples from Skároš, Slovakia.

Fig. 6. Bryozoan Schizoporella cf. tetragona (Reuss, 1847) from Skároš, Slovakia (NHMW 2015/0400/0010). a – cast fossil; b – mold
fossil; c – as previous, colour inverted. All photographs by OM.

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tor  (Adam  Tomašových)  are  thanked  for  constructive  criti-
cism. The senior author (MH) has been supported by Austrian
Science  Fund  (FWF;  Lise  Meitner  Program  M  1544-B25)
and  the  Slovak  Research  and  Development  Agency  under
contracts  no.  APVV-0099-11,  APVV-0436-12,  APVV-SK-
CZ-2013-0129.

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