GeolCarp_Vol66_No3_217

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, JUNE 2015, 66, 3, 217—233 doi: 10.1515/geoca-2015-0021

Introduction

Since the Mesozoic, decapod crustaceans have been increas-
ingly significant components of marine benthic invertebrate
associations  of  the  continental  shelf  and  slope  (Glaessner
1969; Feldmann 2003; Klompmaker et al. 2013; Noël et al.
2014). Especially brachyuran crabs are among the most suc-
cessful  of  all  malacostracan  crustacean  groups  in  terms  of
number  of  ecological  niches  (Warner  1977;  Schram  1986;
Taylor & Schram 1999) and sheer number of species (Ng et
al. 2008; De Grave et al. 2009; Schweitzer et al. 2010). Dur-
ing  the  Miocene,  one  of  the  major  decapod  diversification
events  occurred  (Schweitzer  2001;  Feldmann  &  Schweitzer
2006). In the Western Tethys area this was enhanced by the
biogeographical  differentiation  at  that  time  resulting  in  two
different paleogeographical areas, circum-Mediterranean and
Paratethys  (Rögl  1998,  1999;  Harzhauser  et  al.  2002;
Harzhauser  &  Piller  2007),  and  leading  to  complex  migra-
tion patterns in marine benthic associations (Studencka et al.
1998;  Kroh  &  Harzhauser  1999;  Harzhauser  et  al.  2003,
2007, 2008; Moissette et al. 2006; Kocsis et al. 2012).

In the second half of the 20

century, Miocene decapod

crustacean assemblages of the Western and Central Para-
tethys were studied by Friedrich Bachmayer (Bachmayer

Decapod Crustacea of the Central Paratethyan Ottnangian

Stage (middle Burdigalian): implications for systematics and

biogeography

MATÚŠ HYŽNÝ

1,2!

, MATHIAS HARZHAUSER

and WOLFGANG DANNINGER

Geological-paleontological Department, Natural History Museum Vienna, Burgring 7, 1010 Vienna, Austria;

hyzny.matus@gmail.com; mathias.harzhauser@nhm-wien.ac.at

Department of Geology and Paleontology, Faculty of Natural Sciences, Comenius University, Ilkovičova 6, 842 15 Bratislava,

Slovak Republic

Hauptstrasse 83, 4794 Kopfing, Austria; wolfgang.danninger@gmail.com

(Manuscript received November 4, 2014; accepted in revised form March 12, 2015)

Abstract: Decapod crustaceans from the Ottnangian (middle Burdigalian, Lower Miocene) of the Western and Central
Paratethys remain poorly known. In this study, we review and re-describe mud shrimps (Jaxea kuemeli), ghost shrimps
(Gourretia sp., Calliax michelottii) and brachyuran crabs of the families Leucosiidae, Polybiidae and Portunidae. A
dorsal carapace of the genus Calliax is reported for the first time in the fossil record. Re-examination of the type
material of Randallia strouhali (Leucosiidae) and Geryon ottnangensis (Geryonidae) resulted in a transfer of these
species into Palaeomyra (Leucosiidae) and Liocarcinus (Polybiidae), respectively. Achelous vindobonensis, originally
described as a chela of a portunid crab, probably belongs to a member of Polybiidae and is provisionally treated as
Liocarcinus sp. Only two species, J. kuemeli and C. michelottii, are also known from the Karpatian, the succeeding
Paratethyan stage. In most cases, the decapod assemblages of the Ottnangian consist of rather shallow-water taxa whereas
the assemblages of the Karpatian consist of deep-water taxa from the middle and outer shelf. The Central Paratethyan
assemblages show similarities in genus composition to the Proto-Mediterranean and recent Indo-Pacific regions. Gourretia
sp. represents the earliest occurrence of the respective genus in the fossil record. The Oligocene—Early Miocene appear-
ance of Palaeomyra and Liocarcinus in the circum-Mediterranean implies that sources of present-day diversity hotspots
in the Indo-Pacific trace to the Western Tethys (as for other decapod genera), although coeval decapod assemblages in
the Indo-Pacific remain poorly known.

Key words: Crustacea, Decapoda, Central Paratethys, Ottnangian, Early Miocene.

1950, 1953a,b,c, 1954, 1962, 1982; Bachmayer & Tollmann
1953),  Reinhard  Förster  (Förster  1979a,b)  and  Pál  Müller
(Müller 1984, 1996, 1998a, 2006). Recently a renewed inter-
est  in  these  faunas  provided  new  data  on  Ukraine  (Rad-
wański  et  al.  2006;  Ossó  &  Stalennuy  2011),  Slovenia
(Mikuž  2003,  2010;  Mikuž  &  Pavšič  2003;  Gašparič  &
Hyžný  2014)  and  Slovakia  (Hyžný  2011a,b,c;  Hyžný  &
Schlögl  2011;  Hyžný  &  Hudáčková  2012),  as  well  as  new
insights  into  the  taxonomy  of  Central  Paratethyan  ghost
shrimps (Hyžný 2012; Hyžný & Müller 2010, 2012; Hyžný
& Gašparič 2014; Hyžný & Dulai 2014). However, most of
the  work  has  been  done  on  Badenian  material  (see  Müller
1984  for  an  overview)  because  the  Middle  Miocene  Bade-
nian  ( = Langhian  and  lower  Serravallian)  sediments  are  ex-
posed to a great extent in Austria, Slovakia, Hungary, Poland
and Ukraine (Rasser & Harzhauser 2008). In contrast, only a
few outcrops exposing good sedimentological record of Ott-
nangian  and  Karpatian  deposits  exist.  Müller  (1998a)  de-
scribed  a  small  Karpatian  assemblage  from  the  Korneuburg
Basin  (Austria)  and  recently,  Hyžný  &  Schlögl  (2011)  and
Gašparič  &  Hyžný  (2014)  described  Karpatian  deep-water
(epibathyal)  decapod  crustacean  assemblages  from  the  Slo-
vak  part  of  the  Vienna  Basin  and  the  Slovenian  part  of  the
Styrian Basin. Decapods from the Ottnangian strata were re-

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ported by Bachmayer (1953a, 1982) and briefly summarized
by Müller (1998b).

The present contribution aims: 1) to provide a systematic

overview of all decapods from the Ottnangian stage, includ-
ing the re-examination of older material of Bachmayer
(1953a) and report on previously undescribed specimens in-
cluding new localities with decapod occurrences, and 2) to
discuss the paleobiogeographic affinities of the decapod as-
semblages and implications for their evolutionary patterns.

Geological setting

The material derives largely from the North Alpine Fore-

land Basin (NAFB) and partly from the Vienna Basin (VB).
During the early and middle Ottnangian, the NAFB was part
of a marine gateway, referred to as the Burdigalian Seaway
(Allen et al. 1985). This strait connected the western Proto-
Mediterranean Sea with the Central Paratethys and was char-
acterized by extensive shelf areas bordered by the advancing
Alpine thrust front. Along the northern shelf, widespread tidal
deposits developed under meso- and macrotidal conditions
(Pippèrr 2011; Grunert et al. 2012). At that time, the Vienna
Basin had not formed yet and the area was part of the junc-

tion  between  the  NAFB  and  the  Carpathian  Foreland  Basin
(Kováč  et  al.  2004),  characterized  by  offshore  deposits  of
“Schlier”-type. As a result of the uplift of the NAFB, the ma-
rine connections ceased and fluvial-lacustrine environments
of  the  Upper  Freshwater  Molasse  were  established  in  the
western  NAFB  (Berger  1996;  Reichenbacher  et  al.  2013)
during  the  late  Ottnangian.  No  decapod  remains  are  known
from  that  phase.  The  decapod  crustaceans  reviewed  here
come from several localities (Fig. 1):

Neuhofen bei Tettenweis (Bavaria, Germany) – NAFB,

early  Ottnangian:  This  is  the  type  locality  of  the  Neuhof
Beds  (Neuhofener  Schichten).  It  consists  of  clayey  to  fine-
grained sandy marls (Doppler et al. 2005). The Neuhof Beds
in  the  Eastern  Mollasse  are  the  equivalent  of  the  Kalkofen
Formation in the Western Molasse, and their age is early Ott-
nangian  (Heckeberg  et  al.  2010).  Based  on  foraminifer-as-
semblages,  Pippèrr  (2011)  assumed  deposition  on  the  outer
shelf for the Neuhof Beds.

Ottnang/Schanze (Upper Austria) – NAFB, early Ott-

nangian: This section has been chosen as the stratotype for
the regional Ottnangian stage by Rögl et al. (1973). It is part
of an abandoned clay pit near a memorial to the Peasant
Wars (called “Schanze”) and has been declared a natural her-
itage site. Recently it was revised by Grunert et al. (2010a,

Fig. 1. Geographic position of the studied localities (triangles). Neogene basins in grey (a) and white (b). Maps modified after Grunert et
al. (2010a) and Kroh (2005).

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2012), who proposed an age of 17.95—18.056 Ma for the sec-
tion.  The  section  is  about  10  m-thick  and  exposes  clayey
silts and clayey—sandy silts with sandy lenses and flaser bed-
ding  passing  into  mollusc-rich  and  bioturbated  pelitic  sedi-
ments.  The  analysis  by  Grunert  et  al.  (2012)  revealed  a
eutrophic environment at the transition from a suboxic outer
shelf  to  upper  bathyal  towards  a  better  oxygenated  middle
shelf  environment  under  the  influence  of  storm  events  and
tidal currents.

Allerding (Upper Austria) – NAFB, early Ottnangian:

The Allerding locality is situated in the quarry of the Schär-
dinger Granit Industrie AG close to Schärding in Upper Aus-
tria.  At  this  quarry,  the  Moldanubian  Schärding  Granite  is
exploited. It is part of the NW-SE trending margin of the Bo-
hemian  Massif,  which  also  formed  the  coast  of  the  western
Central  Paratethys  during  most  of  the  Early  Miocene.  The
granite  is  overlain  by  a  few  meters  of  Ottnangian  deposits
(Harzhauser  et  al.  2014).  The  presence  of  coastal  boulders
and  the  mollusc  assemblages  suggest  a  rocky  shore,  which
quickly became sealed by blue-grey pelitic “Schlier” during
a major transgression (Harzhauser et al. 2014).

Grosswiesenhart (Upper Austria) – NAFB, early Ott-

nangian: The material derives from the clays of a claypit at
St. Marienkirchen close to Schärding. Little paleontological
information is available for the section aside from a note on
the  occurrence  of  the  dogfish  shark  Isistius  Gill,  1865  in
Schultz  (2013).  According  to  the  geological  map  (Rupp
2008)  the  section  is  part  of  the  lower  Ottnangian  Ottnang
Formation.  The  depositional  environment  was  most  proba-
bly shelfal but no details are known to us.

Limberg (Lower Austria) – NAFB, early Ottnangian:

The Limberg locality exposes a few meters of finely laminated
diatomites and clays of the Limberg Member at the base of
the pelitic Zogelsdorf Formation. The section was studied by
Roetzel et al. (1996) and Grunert et al. (2010b), who docu-
mented upwelling conditions in a distal-shelf environment
with nutrient-rich waters established during the early Ottnan-
gian along the steep escarpment of the Bohemian Massif in
the eastern NAFB.

Ort im Innkreis (Upper Austria) – NAFB, early Ott-

nangian: The outcrop exposes “Schlier” deposits, namely
clayey silts and clays—sandy silts, which are a part of the lower
Ottnangian Ottnang Formation according to the geological
map (Krenmayr & Schnabel 2006; Rupp 2008). The deposi-
tional environment was most probably shelfal but no details
are known to us.

Peterskirchen (Upper Austria) – NAFB, middle Ott-

nangian: A single specimen in the NHMW collections was
found in one of the clay pits at Peterskirchen, which expose
light-grey to greenish-grey marly clays and silts (Ćorić 1998).
These  deposits  are  part  of  the  middle  Ottnangian  Ried  For-
mation  (Rupp  2008)  and  formed  in  an  inner  shelf  environ-
ment  under  low-energy  with  high  abundance  of  Ammonia
Brünnich, 1772 (Rupp & van Husen 2007).

Antiesen bluff (Upper Austria) – NAFB, middle Ott-

nangian: The Ottnang Formation is overlain by mica-rich
sand with cross bedding and pelitic intercalations of the mid-
dle Ottnangian Reith Formation (Rupp 2008). These are well
exposed along the banks of the Antiesen river, 1.7 km SE of

Antiesenhofen. The decapods were collected from these pel-
itic intercalations. The microfauna is dominated by Ammonia
and the depositional environment was interpreted by Rupp &
van Husen (2007) and Rupp (2008) as a tidal-influenced in-
ner shelf setting.

Pramhof at Schärding (Upper Austria) – NAFB, early

or  middle  Ottnangian:  Little  information  is  available  for
this locality. According to the geological map (Rupp 2008),
the  section  is  part  of  the  lower  Ottnangian  Ottnang  Forma-
tion. C. Rupp (pers. comm.), however, pointed out that rem-
nants of the middle Ottnangian Ried Formation also exist in
the  area.  Decapod  specimens  are  preserved  in  the  pelitic
“Schlier”, however, it cannot be decided from which forma-
tion they actually come.

Grosskrut 3 (Lower Austria) – VB, middle Ottnan-

gian: This is the only locality in the Vienna Basin – the ma-
terial  derives  from  a  core  (depth  500 m),  which  drilled  into
the  Ottnangian  Lužice  Formation.  The  sample  with  a  deca-
pod  corresponds  to  the  typical  lithology  of  the  Lužice  For-
mation,  with  laminated  grey  calcareous  clays,  silt  and
siltstones  with  intercalations  of  sands.  The  decapod  sample
was  collected  from  silty  clay.  The  Lužice  Formation  indi-
cates  open  marine  shelf  settings  with  widespread  dysoxic
bottom conditions (Kováč et al. 2004).

Material and methods

Studied material comes either from older collections

(GBA, KM, NHMW) or was newly collected by one of us
(WD). Preparation, if needed, was done with a pneumatic
needle. Specimens were photographed either dry and uncoated
or covered with ammonium chloride sublimate.

The repositories of specimens illustrated or referred to be-

low are as follows:

GBA – Geological Survey, Vienna (Austria)
NHMW – Natural History Museum, Vienna (Austria)
KM – Krahuletz-Museum, Eggenburg (Austria).

Systematic paleontology

Order: Decapoda Latreille, 1802

Infraorder: Gebiidea de Saint Laurent, 1979

Family: Laomediidae Borradaile, 1903

Genus Jaxea Nardo, 1847

Type species: Jaxea nocturna Nardo, 1847, by original

designation.

Diagnosis: See Ngoc-Ho (2003; p. 501).
Remarks: The fossil record of the genus has recently

been  revisited  by  Hyžný  (2011c).  He  showed  that  the  tooth
formula and general cheliped shape is more-or-less constant
and  on  its  basis  the  identification  to  species  level  can  suc-
cessfully be made. Two extant species, Jaxea nocturna Nardo,
1847  from  the  Mediterranean  and  adjacent  European  seas
(Ngoc-Ho 2003) and J. novaezealandiae Wear & Yaldwyn,
1966  from  New  Zealand  (Wear  &  Yaldwyn  1966),  differ
markedly  in  their  tooth  formula.  The  only  fossil  species,

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J. kuemeli Bachmayer, 1954 appears to be much closer to the
Mediterranean species (Hyžný 2011c).

Jaxea kuemeli Bachmayer, 1954

Figs. 2—3

v*1954 Jaxea kümeli Bachmayer, 1954, p. 64, pl. 1, figs. 1, 2
1969    Jaxea kuemeli Bachmayer – Glaessner, p. R477, fig. 284.4a, 4b
1984    Jaxea kuemeli Bachmayer – Müller, p. 49
1998    Jaxea kümeli Bachmayer – Mayoral et al., p. 508
2011c  Jaxea  kuemeli  Bachmayer  –  Hyžný,  p. 176  figs. 2A—D,  3B,

4A—C, 5A—C, 6A—D

2014 Jaxea kuemeli Bachmayer – Gašparič & Hyžný, p. 9, figs. 8—9
(for comprehensive synonymy see Hyžný 2011c, p. 176).

Diagnosis: See Hyžný (2011c; p. 176)
Material examined: Ottnang: 19 specimens (under col-

lective number GBA 2009/014/023), two near-complete spec-
imens  (NHMW  2009/0150/0001,  NHMW  2010/0090/0001);
Pramhof  at  Schärding:  one  specimen  retaining  both  cheli-
peds (NHMW 2009/0160/0001); Grosswiesenhart: one spec-
imen  retaining  both  chelipeds  (NHMW  1986/0101/0023).
Some  specimens,  specifically  those  from  Ottnang,  possess
cuticular surfaces.

Discussion: The studied material fully corresponds to

Jaxea kuemeli as re-described by Hyžný (2011c). For the

Fig. 2. Jaxea kuemeli Bachmayer, 1954. a—b – near-complete specimen from Ottnang (NHMW 2009/0150/0001), c—d – isolated cheli-
peds from Grosswiesenhart (NHMW 1986/0101/0023). Scale bar equals 5 mm.

species, the most obvious distinctive character is the position
of the large median tooth on the cutting edge of the pollex.
In J. kuemeli this is located posteriorly relative to the posi-
tion of the median tooth on the dactylus (Hyžný 2011c;
fig. 3B), while it is positioned anteriorly in its close relative,
the extant J. nocturna (Hyžný 2011c; fig. 3A).

Occurrence: The species has been identified in lower

and middle Miocene (upper Burdigalian to Langhian) strata
of Austria, Slovenia, Slovakia and Hungary (Bachmayer
1954; Hyžný 2011c; Gašparič & Hyžný 2014).

Infraorder: Axiidea de Saint Laurent, 1979

Family: Ctenochelidae Manning & Felder, 1991

Genus: Gourretia de Saint Laurent, 1973

Type species: Callianassa denticulata Lutze, 1937 ( = Cal-

lianassa subterranea var. minor Gourret, 1887) by original
designation.

Diagnosis: See Ngoc-Ho (2003; p. 498).
Remarks: The fossil record of the genus is poorly

known. Vega et al. (2007) re-assigned Callianassa aquilae
Rathbun, 1935 from the Upper Cretaceous strata of Texas,
Louisiana and Mexico to Gourretia de Saint Laurent, 1973.
Recently, Hyžný in Bermúdez et al. (2013) erected a new ge-
nus, Rathbunassa, for the species, leaving Gourretia without

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fossil  record.  Material  from  Ottnang  exhibits  striking  simi-
larity  to  the  major  propodus  of  Callianassa  denticulata
Lutze,  1937,  the  type  species  of  Gourretia  (compare  with
López  de  la  Rosa  et  al.  1998,  fig. 1B;  Ngoc-Ho  2003,
fig. 21L; Sakai & Türkay 2005, fig. 2A). It is assigned to this
genus  based  on  a  number  of  characters:  upper  margin  is
slightly  converging  distally,  the  fingers  are  shorter  than
manus (palm), dactylus has no prominent teeth on its occlusal
margin  and  is  hooked  distally.  These  features  alone  can
hardly be considered of taxonomic importance, however, their
combination  has  affinities  to  Gourretia.  The  generic  assign-
ment,  though,  should  be  considered  preliminary  until  more
complete  material  is  revealed.  If  confirmed  its  attribution  to
Gourretia,  the  occurrence  from  Ottnang  discussed  below  is
the only known report of the genus in the fossil record.

Gourretia sp.

Fig. 4

Fig. 3. Jaxea kuemeli Bachmayer, 1954. a – near-complete specimen (NHMW 2010/0090/0001), b – near-complete specimen (GBA
2009/014/023 coll. number), c—d – isolated chelae (GBA 2009/014/023 coll. number). All specimens are from Ottnang and were covered
with ammonium chloride prior to photography. Scale bar equals 5 mm.

Fig. 4. Gourretia sp. from Ottnang (GBA 2009/014/0024). a – covered with ammonium chloride, b – uncoated, c – reconstruction of
the chela outline. Scale bar equals 5 mm.

Material examined: Ottnang: major left propodus artic-

ulated with dactylus plus two isolated dactyli, all with pre-
served cuticle (collective number GBA 2009/014/0024).

Measurements: Propodus max. length incl. fixed

finger = 25 mm; propodus max height = 13 mm; propodus
min. height = 11.2 mm; dactylus length = 9 mm (measure-
ments taken on articulated specimen).

Description: Propodus longer than high, converging dis-

tally; upper and lower margins smooth, without any apparent
serration; fixed finger two times shorter than manus, occlusal
surface with one prominent tooth at its base; tip of the fixed
finger pointed and bent slightly upward; dactylus robust and
edentulous.

Discussion: Many Miocene decapod species that are

congeneric with the extant ones represent distinct taxa (e.g.
Müller 1984; Hyžný 2011c; Hyžný & Müller 2012), thus it is
fairly possible that the material studied here represents a new
species. Propodus and dactylus, however, cannot be consid-

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ered of taxonomic importance as these elements are often sub-
ject to intraspecific variation. The studied specimens are treated
in open nomenclature until more complete material is found.

Occurrence: The species is restricted to the Lower Ott-

nangian of Ottnang beds, Austria.

Family: Callianassidae Dana, 1852

Subfamily: Eucalliacinae Manning & Felder, 1991

Genus Calliax de Saint Laurent, 1973

Type species: Callianassa (Callichirus) lobata de Gail-

lande & Lagardère, 1966 by original designation.

Diagnosis: See Hyžný & Gašparič (2014; p. 42).
Remarks: Calliax de Saint Laurent, 1973 has a complex

taxonomic history. It has been discussed in detail by Hyžný
(2012) and Hyžný & Gašparič (2014) and there is no need to
repeat it here. Discussion on distinguishing Calliax from re-
lated  taxa  based  on  soft-part  morphology  was  provided  by
Ngoc-Ho  (2003).  Recently,  Hyžný  &  Gašparič  (2014)  dis-
cussed  in  detail  the  identification  of  the  genus  in  the  fossil
record:  rectangular  major  cheliped  propodus  with  two  ridges
on  the  base  of  the  fixed  finger  extending  onto  manus  with
combination of relatively short fingers is typical for the genus.

Calliax michelottii (A. Milne-Edwards, 1860)

Fig. 5

1860 Callianassa Michelotti n. sp. – A. Milne-Edwards, p. 341, pl. 14,

fig. 3

1928 Callianassa Michelottii A. Milne-Edwards, 1860 – Glaessner,

p. 167—168

1984 Callianassa szobensis n. sp. – Müller, p. 53, pl. 7, figs. 3—4
2014 Calliax michelottii Hyžný & Gašparič, p. 45, figs. 5—10
2014 Calliax michelottii Gašparič & Hyžný, p. 5, figs. 4—5
(for comprehensive synonymy see Hyžný & Gašparič 2014, p. 45).

Diagnosis: See Hyžný & Gašparič (2014; p. 45).
Material examined: Neuhofen bei Tettenweis: left ma-

jor chela (NHMW 2010/0089/0001); Peterskirchen: major
left chela associated with dorsal carapace consisting of part
and counterpart (NHMW 2014/0412/0001); Ort: major right
propodus associated with carpus (NHMW 2014/0405/0001);

Antiesen: isolated major left propodus (NHMW 1985/0067/
0026). Only specimens from Ort and Antiesen retain cuticu-
lar surfaces.

Description: Carapace without dorsal oval and with car-

diac prominence; major cheliped merus longer than high and
keeled  along  its  midline,  upper  margin  straight,  lower  mar-
gin poorly preserved; carpus higher than long, upper margin
concave, proximo-lower margin rounded; propodus distinctly
longer  than  high,  upper  and  lower  margins  parallel  to  each
other;  fixed  finger  with  double  ridge  running  onto  manus;
outer lateral surface of propodus covered with several tuber-
cles near the base of the fixed finger; dactylus deep and ro-
bust;  occlusal  surface  of  both  fingers  armed  with  a  row  of
tiny teeth.

Discussion: The species was originally described as Cal-

lianassa  Leach,  1814.  Recently,  Hyžný  &  Gašparič  (2014)
redescribed  the  species  and  transferred  it  to  Calliax.  The
re-description  is  based  on  the  chelae,  as  these  are  often  the
only  fossil  remains  known  from  ghost  shrimps  (Bishop  &
Williams  2005).  In  this  respect,  the  material  from  Peters-
kirchen is exceptional because it also exhibits features of the
dorsal  carapace  which  is  usually  not  preserved  in  ghost
shrimps because of its weak calcification (but see e.g. Hyžný
& Schlögl 2011, text-fig. 2C for the opposite). Dorsal cara-
pace of the Peterskirchen specimen possesses cardiac promi-
nence which is also present in extant Calliax doerjesti Sakai,
1999.  Features  of  the  major  chela  fully  conform  to  the  de-
scription made by Hyžný & Gašparič (2014).

Occurrence: Late Oligocene—Middle Miocene of Ger-

many, Italy, Austria, Slovenia, Slovakia and Hungary
(Hyžný & Gašparič 2014; Gašparič & Hyžný 2014).

Infraorder: Brachyura Linnaeus, 1758

Section: Eubrachyura de Saint Laurent, 1980

Subsection: Heterotremata Guinot, 1977

Superfamily: Leucosioidea Samouelle, 1819

Family: Leucosiidae Samouelle, 1819

Subfamily: Ebaliinae Stimpson, 1871

Genus Palaeomyra A. Milne Edwards in Sismonda, 1861

= Ebalites Müller, 1978

? = Tanaoa Galil, 2003

Fig. 5. Calliax michelottii (A. Milne-Edwards, 1860). a—c – specimen from Peterskirchen (NHMW 2014/0412/0001) in moulting position
with preserved chelae and dorsal carapace; interpretative drawing in (b). d – isolated carpus and propodus from Ort (NHMW 2014/0405/0001).
Scale bar equals 5 mm.

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Type species: Palaeomyra bispinosa A. Milne Edwards

in Sismonda, 1861 by original designation and monotypy.

Diagnosis: Carapace subcircular, globose, covered with

granules.  Front  narrow,  bilobed.  Outer  orbital  margin  with
three  sutures.  Branchial  and  intestinal  regions  swollen,  de-
marcated  by  grooves,  2  pairs  of  pits  along  cardiobranchial
grooves. Posterior margin narrow and bidentate.

Remarks: Palaeomyra was erected by A. Milne Edwards

in Sismonda (1861) to accommodate a single species, P. bis-
pinosa A. Milne Edwards in Sismonda, 1861 from the Mio-
cene of Torino, Italy. Although the original material, a single
specimen (holotype), has been lost (De Angeli & Garassino
2006), Garassino et al. (2004) reported six additional and
more complete specimens from the Oligocene of Morbello,
Alessandria and the Upper Miocene of Cocconato, Asti, both
in Italy, which helped the re-evaluation of the species.

Müller (1978) erected Ebalites for Ebalia globulosa

Müller,  1975  from  the  Middle  Miocene  of  Hungary.  Later,
he  (Müller  1984)  recognized  the  genus  to  be  synonymous
with  Palaeomyra  and  reassigned  E.  globulosa  once  more,
this  time  to  Palaeomyra.  Until  now,  P.  bisponosa  and  P.
globulosa  have  been  considered  the  only  known  species  of
the genus (Schweitzer et al. 2010). Based on the major simi-
larities in the carapace outline, swollen branchial and intesti-
nal  regions  demarcated  by  grooves,  2  pairs  of  pits  along
cardiobranchial grooves, posterior margin with 2 prominent
spines  and  distinct  granulation  covering  the  surface  of  the
carapace, the specimen from Ottnang, originally described as
Randallia  strouhali  Bachmayer,  1953a  is  transferred  here
into Palaeomyra.

Since no generic diagnosis of Palaeomyra was given by

Glaessner (1969) or any subsequent author, it is provided
here based on the description and figure of A. Milne Ed-
wards in Sismonda (1861; p. 14, figs. 18—20) and additional
material reported by Garassino et al. (2004).

Galil (2003) revised selected species of Randallia Stimp-

son, 1857 and erected four new genera, including Tanaoa
Galil, 2003. Interestingly, Bachmayer (1953a) noted similar-
ities between Randallia strouhali and extant R. pustulosa

Wood-Mason  in  Wood-Mason  &  Alcock,  1891,  the  type
species  of  Tanaoa.  The  dorsal  carapace  morphology  of
Palaeomyra  (Müller  1984;  pl. 47,  figs. 1—6)  corresponds  to
the variation of Tanaoa as expressed in Galil (2003; fig. 1B—D).
In fact, swollen branchial and intestinal regions demarcated
by grooves and 2 pairs of pits along cardiobranchial grooves
are  the  characters  typical  nor  only  for  Palaeomyra  but  also
for  Tanaoa  and  Toru  Galil,  2003  (Galil  2003),  whereas
strong  granulation  on  the  entire  surface  of  the  dorsal  cara-
pace  of  Palaeomyra  is  closer  to  Tanaoa.  These  two  genera
may well be considered synonymous, but we are reluctant to
synoymize them without direct comparison with some extant
material of Tanaoa.

Based on the supposed close relationship between Palaeo-

myra and Tanaoa, former taxon, previously unplaced at sub-
family level, they are assigned here to the subfamily
Ebaliinae.

Palaeomyra strouhali (Bachmayer, 1953a) n. comb.

Fig. 6

1953a Randallia strouhali Bachmayer, p. 137, pl. 5, figs. 1—2
1998b Randallia strouhali Bachmayer – Müller, p. 24, pl. 2, fig. 1

Emended diagnosis: Almost circular carapace with

nearly straight posterolateral margins and bidentate posterior
margin; dorsal surface covered with densely packed granules
of two sizes; cardiac and intestinal regions without any tu-
bercles or spines.

Material examined: Holotype NHMW 1953/0051/0001

showing incomplete dorsal carapace with preserved cuticle.

Measurements: Maximum length of the fragmentary

carapace = 7 mm; greatest width = 8.5 mm.

Emended description: Carapace almost circular in out-

line, widest approximately at midlength; anterior portion not
preserved.  Posterolateral  margins  nearly  straight.  Posterior
carapace  margin  straight  and  narrow,  with  two  stubby  pro-
trusions.  Dorsal  surface  of  carapace  covered  with  many
large,  densely  packed  granules,  nearly  identical  in  size  (dia-

Fig. 6. Palaeomyra strouhali (Bachmayer, 1953a). Holotype (NHMW 1953/0051/0001) from Ottnang. a – detail of the posterior portion
of the carapace; note the tuberculation, b – entire specimen, whitened with ammonium chloride prior to photography, c – interpretative
drawing with carapace regions indicated.

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meter  0.3 mm),  interspaced  with  smaller  ones  (diameter
0.1 mm).  Two  pairs  of  pits  present  along  cardiobranchial
grooves.  Hepatic,  gastric,  cardiac  and  branchial  regions  de-
marcated  with  grooves;  grooves  deepest  in  posterior  cara-
pace  portion.  Gastric  region  large  and  only  indistinctly
demarcated with grooves. Cardiac region circular in outline
and strongly arched. Branchial regions broad and ovate. In-
testinal region narrow, without any tubercles.

Discussion: Bachmayer (1953a) described the species

based on a single incomplete specimen. The description is
emended here as a result of re-examination of the holotype;
some characters (pits along cardiobranchial grooves) were
omitted in the original description.

The most-anterior portion of the holotype is missing, so the

frontal margin cannot be described. Nevertheless, the cuticular
surface and distinct groove pattern in the posterior portion of
the carapace allow distinguishing of P. strouhali from its two
congeners. Garassino et al. (2004; p. 267) noted the variation
in dorsal tuberculation of P. bispinosa and the same is appar-
ent from the figures of P. globulosa published by Müller
(1984; pl. 47, figs. 3—6). However, none of these species has
tubercles as flattened and densely packed as P. strouhali.

Palaeomyra strouhali also differs from six extant species of

Tanaoa. Most of the extant species, including Tanaoa distinc-
tus (Rathbun, 1894), T. nanus Galil, 2003, T. pustulosus and

T. serenei (Richer de Forges, 1983) possess a cardiac spine
(Galil 2003; Ng & Richer de Forges 2007), which is lacking in
P. strouhali. Granulation of the carapace seems to be distinct
in different species of Tanaoa. In this respect, P. strouhali is
closest to T. serenei (Ng & Richer de Forges 2007, fig. 1D).

Occurrence: The species is known only from the early

Ottnangian of Ottnang, Austria.

Superfamily: Portunoidea Rafinesque, 1815

Family: Polybiidae Ortmann, 1893

Genus: Liocarcinus Stimpson, 1871

Type species: Portunus holsatus Fabricius, 1798, by

original designation.

Diagnosis: See Manning & Holthuis (1991; p. 83).
Remarks: Recent analyses (Schubart & Reuschel 2009;

Spiridonov  et  al.  2014)  resolved  the  paraphyletic  nature  of
the  genus.  Revision  of  all  extant  species  is  currently  in
progress  (Schubart  &  Reuschel  2009).  Considering  rather
high morphological differences among fossil taxa attributed
to Liocarcinus so far (Schweitzer et al. 2010), the revision of
the fossil record of the genus is needed as well.

Liocarcinus ottnangensis (Bachmayer, 1953a) n. comb.

Figs. 7—9

Fig. 7. Liocarcinus ottnangensis (Bachmayer, 1953a). a, d – holotype (NHMW 1953/0052/0001) from Ottnang, b, c, f – specimen from
Allerding (NHMW 2014/0402/0001), part (b) and counterpart (c), e – line drawing of the holotype showing the number of frontal and
anterolateral teeth, f – detail of the anterolateral margin with four anterolateral teeth (empty arrows) and outer orbital tooth (filled arrow).
Specimens in a—c were covered with ammonium chloride prior to photography. Scale bar equals 5 mm.

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Fig. 8. Liocarcinus ottnangensis (Bachmayer, 1953a). a—c – female specimen in ventral aspect from Pramhof (NHMW 1971/1486 A),
b—c – detail of the sternum with indicated press-button structures (filled arrows) and vulvae (empty arrows). Scale bar equals 5 mm.

Fig. 9. Liocarcinus ottnangensis (Bachmayer, 1953a). a—b – frag-
mentary fixed finger (NHMW 2014/0402/0004) in occlusal and lat-
eral  view,  c—d  –  fragmentary  fixed  finger  (NHMW  2014/0402/
0005)  in  occlusal  and  lateral  view,  e  –  fragmentary  dactylus
(NHMW  2014/0402/0006)  in  lateral  view,  f  –  fragmentary  fixed
finger (NHMW 2014/0402/0007) in lateral view. All specimens are
from Allerding. Scale bar equals 5 mm.

1953a Geryon ottnangensis Bachmayer, p. 138, pl. 5, figs. 3—4
1998b Geryon ottnangensis Bachmayer – Müller, p. 37
2010 Chaceon ottnangensis (Bachmayer); Schweitzer et al., p. 106

Emended diagnosis: Carapace hexagonal, 1.1 times

wider than long; front with three prominent teeth, orbits deep
and rounded; anterolateral margin with four large teeth (ex-
cluding outer orbital tooth), third smallest but distinct; first
three anterolateral teeth equally spaced from each other, dis-
tance between the third and fourth somewhat greater. Cardiac
region wide and elevated, bearing two swellings at widest
points; epibranchial regions with prominent transverse ridge
running across the mesogastric region.

Material examined: Ottnang: holotype of Geryon ott-

nangensis  NHMW  1953/0052/0001;  Allerding:  dorsal  cara-
pace  NHMW  2014/0402/0001,  fragmentary  cheliped
NHMW  2014/0402/0003,  sixteen  cheliped  fragments
NHMW  2014/0402/0004—0007,  2014/0402/0008  (collective
number);  Pramhof  at  Schärding:  carapace  with  chelipeds  in
ventral aspect exposing female sternum NHMW 1971/1486 A.
Cuticle is entirely or partly preserved in all studied specimens.

Measurements: Holotype NHMW 1953/0052/0001: max.

carapace width = 32.5 mm, max. carapace length = 30.4 mm;
NHMW 2014/0402/0001: max. carapace width = 31.5 mm,
max. carapace length = 29.0 mm (preserved part).

Emended description: Carapace hexagonal, 1.1 times

wider than long, strongly convex from front to posterior

margin;  carapace  surface  covered  with  very  fine  granules
and  pits.  Frontal  margin  with  three  prominent  teeth,  orbits
relatively  deep  and  rounded;  anterolateral  margin  with  four
large  teeth  (excluding  outer  orbital  tooth),  all  sharp,  third
smallest  but  distinct;  first  three  anterolateral  teeth  equally
spaced  from  each  other,  distance  between  the  third  and
fourth somewhat greater. Regions well developed; mesogas-
tric  region  with  a  long  anterior  process  ending  in  a  frontal
sulcus, urogastric region depressed; cardiac region wide and

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elevated, bearing two swellings at widest points; epibranchial
regions with prominent transverse ridge running across the
mesogastric region; branchial regions swollen, with prominent
ridge longitudinally along the midline of the branchial region.

Female sternum ovoid, slightly longer than wide, greatest

width  at  sternite 6,  lateral  margins  forming  a  series  of  arcs
defined by smoothly curved lateral somite margins and nar-
row  episternal  projections;  sternites 1—4  (supposedly)  fused
completely,  elements 1—3  not  preserved.  Axial  depression
prominent, narrow. Vulvae positioned near the anterior mar-
gin  of  sternite 6,  the  genital  opening  ovate  and  slightly  ob-
lique; press-button structure positioned on sternite 5.

First pereiopods (chelipeds) strong. Both fingers bearing

dome-shaped denticles on occlusal surface.

Discussion: The original description of Geryon ottnan-

gensis is based on a single carapace. Bachmayer (1953a; pl. 5,
fig. 4) interpreted the frontal margin as having a single tooth.
Thorough  re-examination  of  the  holotype  revealed  the  pres-
ence of three teeth (excluding inner oribital teeth), a character
typical  of  Liocarcinus  and  Benthochascon  Alcock  &  Ander-
son, 1899. Anterolateral margin of Geryon ottnangensis pos-
sesses  five  teeth  (including  outer  orbital  tooth),  a  character
typical for Liocarcinus (Manning & Holthuis 1981; Froglia &
Manning 1982) whereas Benthochascon has only four antero-
lateral teeth (including outer orbital tooth) (Ng 2000). As a re-
sult,  Geryon  ottnangensis  is  transferred  to  Liocarcinus  here.
Interestingly,  all  five  teeth  can  be  distinguished  on  the  holo-
type,  although  Bachmayer  (1953a)  did  not  recognize  their
number.  Geryon  ottnangensis  Bachmayer,  1953a  is  listed  as
Chaceon  Manning  &  Holthuis,  1989  by  Schweitzer  et  al.

(2010), although no arguments for this act were stated. Cha-
ceon, however, always has four frontal teeth (Manning &
Holthuis 1989; Schweitzer & Feldmann 2000), which is in
contrast with the three frontal teeth of Geryon ottnangensis.

Several Liocarcinus species are known from the Miocene

Paratethys; all of them differ from L. ottnangensis. Liocarci-
nus praearcuatus Müller, 1996 and L. rakosensis (Lőrenthey
in Lőrenthey & Beurlen, 1929) from the Middle Miocene of
Poland  and  Hungary  (Müller  1984,  1996)  have  complete
frontal margins; L. oroszyi Bachmayer, 1953b and L. kuehni
Bachmayer,  1953b  from  the  Middle  Miocene  of  Austria
(Bachmayer  1953b;  Müller  1984,  1998b)  have strongly ele-
vated  branchial  regions  and  distinctly  ornamented  carapaces,
respectively. Liocarcinus lancetidactylus Smirnov, 1929 from
the Lower Miocene of Caucasus has three anterolateral teeth
according to Garassino & Novati (2001), but based on their
figures  the  real  number  may  be  higher.  The  preservation
(flattened  crab  bodies  within  shales)  does  not  allow  further
comparison with L. ottnangensis. Liocarcinus oligocaenicus
Paucă,  1929  from  the  Oligocene  of  Romania  (Paucă  1929),
Czech Republic (Jaroš 1939) and Poland (Jerzmańska 1967)
is preserved in a similar manner and the character of the car-
apace is obscured.

From the extant congeners, L. ottnangensis is closest to

the  type  species  L.  holsatus  from  Europe  (Türkay  2001),
which  differs  in  the  nature  of  its  anterolateral  teeth:  in  L.
ottnangensis  the  distance  between  the  fourth  and  fifth  teeth
is proportionately greater than in L. holsatus.

Occurrence: The species is known from the early to

middle Ottnangian in the North Alpine Foreland Basin.

Fig. 10. Liocarcinus sp. a—b – holotype of Achelous vindobonensis Bachmayer, 1950 (NHMW 2014/0409/0001) from Grosskrut 3,
c—d – fragmentary chelae from Ottnang (GBA 2009/014/0025 coll. number). Specimens in b and c were coated with ammonium chloride
prior to photography. Scale bar equals 5 mm.

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Liocarcinus sp.

Fig. 10

1950 Achelous vindobonensis Bachmayer, p. 137, pl. 1, figs. 5, 5a,b
1998b Portunidae ind. sp. – Müller, p. 32

Material examined: Ottnang: several cheliped frag-

ments GBA 2009/014/0025; Grosskrut 3: holotype of Ache-
lous vindobonensis NHMW 2014/0409/0001, with preserved
cuticle.

Discussion: Bachmayer (1950) described Achelous vin-

dobonensis based on a single incomplete chela coming from
the  Grosskrut 3  drilling.  The  material  was  later  treated  as
Portunidae indet. by Müller (1998b). Dentition of chelipeds
has  been  considered  of  taxonomic  significance  recently
(Spiridonov  et  al.  2014).  The  holotype  of  A.  vindobonensis
possesses  fingers  with  bi-  and  trilobed  serial  conical  teeth.
Comparison  with  chelipeds  of  extant  portunoid  crabs  has
shown close affinities to Liocarcinus (Spiridonov et al. 2014;
fig. 3D), but identification on the species-level is not possi-
ble. We refrain from referring this taxon to nomen dubium so
that the name remains available in case better-preserved ma-

Fig. 11. Necronectinae gen. et spec. indet. a—c – incomplete chela
(NHMW 2014/0402/0002) in lateral (a), mesial (b) and dorso-lateral
(c) aspects. Scale bar equals 5 mm.

terial becomes available which may justify or refute this
species name.

Family: Portunidae Rafinesque, 1815

Subfamily: Necronectinae Glaessner, 1928

Remarks: Glaessner (1928) created a new family Ne-

cronectidae to accomodate  an extinct genus Necronectes
A. Milne-Edwards, 1881 ( = Gatunia Rathbun, 1919). The
phylogenetic  analysis  of  Karasawa  et  al.  (2008a)  recog-
nized  a  clade  grouping  Necronectes,  Scylla  de  Haan,
1833  and  Sanquerus  Manning,  1989  within  the  family
Portunidae. As a result, they treated Necronectinae as the
subfamily-level group and it also appeared as such in the
classifications  of  De  Grave  et  al.  (2009)  and  Schweitzer
et al. (2010), which are followed here. The classification
of  Brachyura  presented  by  Ng  et  al.  (2008)  recognized
extant  genera  Scylla  and  Sanquerus  as  members  of  Por-
tuninae.

Necronectinae gen. et spec. indet.

Fig. 11

Material examined: Allerding: left propodus articu-

lated with dactylus NHMW 2014/0042/0002.

Description: First pereiopods (chelipeds) strong.

Fixed  finger  of  left  chela  straight  with  grooves  running
along  its  entire  length  on  both  lateral  surfaces.  Dactylus
slightly  curved,  elongate.  Both  fingers  bearing  biserial
conical teeth on occlusal surface; left dactylus also bear-
ing large molariform tooth proximally.

Discussion: The studied chela bears strong resem-

blance to claws of Scylla as documented in detail by
Keenan et al. (1998). We are, however, hesitant to assign
the material to this genus, as there is another member of
the subfamily known from the Miocene of Europe, namely
Necronectes (Glaessner 1928, 1933). Therefore, we retain

the fragmentary chela from Allerding in the open nomencla-
ture until more material is recovered.

Portunoidea gen. et sp. indet.

Fig. 12

1982 ?Geryon spec. – Bachmayer, p. 22

1998b ?Geryon sp. – Müller, p. 38

Material examined: Limberg: two dorsoventrally near-

complete flattened specimens deposited in KM (without re-
pository number), without preserved cuticular surfaces.

Discussion: Bachmayer (1982) reported two near-com-

plete  specimens  of  ?Geryon  sp.  preserved  in  diatomites  of
Limberg.  Due  to  the  extreme  flattening  their  preservation
was  not  sufficient  for  closer  identification.  Without  a  well
preserved carapace, no closer identification than Portunoidea
gen. et spec. indet. is possible. Unfortunately, the specimens
reported  and  figured  by  Bachmayer  (1982)  could  not  be
traced in the NHMW collections where they were supposed
to be deposited. Two other specimens from the same locality
and  presumably  of  the  same  taxon  are  deposited  in  the

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Krahuletz-Museum in Eggenburg; the opportunity is taken to
publish photos of these specimens (Fig. 12). The preserva-
tion hinders description of any details.

Discussion

Environmental distribution: During the early and

middle  Ottnangian  the  Paratethys  Sea  in  the  North  Alpine
Foreland Basin was a shelfal sea with strong tidal currents and
distinct upwelling areas (Rupp & van Husen 2007; Grunert et
al. 2010b, 2012). The various sections described in the Geo-
logical settings chapter represent depositional environments of
the  upper  bathyal  to  outer  shelf  (e.g.  Ottnang  Schanze,  Lim-
berg)  to  high  energy  (e.g.  Peterskirchen)  and  protected  (e.g.
Antiesen) inner shelf  settings. For many localities, however,
no detailed paleoecological analysis is available.

Decapod crustaceans may have a rather broad bathymetric

distribution on the genus or family level (Abelló et al. 1988;
McLay 1999; Ingle & Christiansen 2004; Ate„ et al. 2006).
However,  certain  species  prefer  specific  conditions  and  in
some cases can be indicative of unique environmental condi-
tions.  For  example,  many  species  inhabiting  coral  reefs  do
not  occur  in  deeper  settings  and  vice  versa:  crabs  living  on
muddy substrates usually do not occur in reefal environment
(Abele 1976; Poupin & Juncker 2010). This general rule has
also  been  demonstrated  for  Badenian  decapods  of  Central
Paratethys (Müller 1984). There is also a general increase in
bathymetric range towards greater depth; since decapod spe-
cies within the first 50 m display a median range of approxi-
mately  150 m,  whereas  abyssal  faunas  tend  to  have  far
greater ranges (Rosa et al. 2012). This may be connected to
seasonal bathymetric differences in bathyal species reaching
500 m (Mura & Cau 1994).

Two taxa are relatively common in the Ottnangian depos-

its,  namely  Jaxea  and  portunoid  crabs.  Portunoids  have
broad  bathymetric  distribution;  however,  Liocarcinus  is
usually  present  at  depths  of  less  than  100 m  (Manning  &
Holthuis  1981).  The  paleobathymetric  distribution  of  mud
and ghost shrimps Jaxea, Gourretia and Calliax is less clear.
They were reported from a broad range of depths, from the
shallow  settings  to  those  exceeding  100 m  (Ngoc-Ho  2003;

Hyžný 2011c; Hyžný & Gašparič 2014), and even up to
800 m in the case of Calliax (Taviani et al. 2013). Similarly,
Tanaoa (as a possible synonym of Palaeomyra) has been re-
ported from depths of a few tens of meters but also from
depths exceeding 1—200 m (Galil 2003).

At Ottnang/Schanze, the environment has been interpreted

as  transition  from  outer  neritic  to  upper  bathyal  based  on  a
detailed statistical analysis of the microfauna. This is domi-
nated  by  benthic  foraminiferal  assemblages  with  Laeviden-
talina  spp.,  Oridorsalisum  bonatus,  Gyroidinoides  spp.  and
Valvulineria  complanata,  characteristic  for  an  outer  neritic
to bathyal setting (Grunert et al. 2012 and references there-
in). Numerous individuals of Jaxea kuemeli that occur at this
locality, frequently with articulated carapace and pleon, sug-
gest  minor  post-mortem  transport.  The  influence  of  occa-
sional storm events and tidal currents in the upper part of the
succession, which formed in inner shelf settings (Grunert et
al. 2012), is a possible cause why brachyurans at this locality
are  known  only  from  isolated  carapaces  (Palaeomyra,  Lio-
carcinus),  whereas  preservation  of  articulated  specimens  of
Jaxea was most probably promoted by the burrows, in which
the  animals  live  (Pervesler  &  Dworschak  1985).  The  near-
complete  body  fossils  of  J.  kuemeli  from  Pramhof  and
Grosswiesenhart may, thus, indicate that they were actually
preserved within their burrows.

At Peterskirchen, the Ottnangian deposits were formed in a

shallow marine low energy environment with high amounts of
Ammonia (Rupp & van Husen 2007). The specimen of Calliax
michelottii  with  both  articulated  chelipeds,  including  cara-
pace,  is  in  accordance  with  this  interpretation  of  low-energy
conditions  and  suggests  autochthonous  preservation  coupled
with rapid burial.

At Allerding, coastal boulders and the mollusc assemblages

indicate that a rocky shore environment was sealed by pelitic
“Schlier” during a transgression event (Harzhauser et al.
2014). The presence of Liocarcinus can be connected to the
transgressive event of the “Schlier” deposition, as this genus
prefers muddy substrates (Rufino et al. 2005).

At Antiesen bluff a tidal-influenced shelf setting was

present (Rupp & van Husen 2007; Rupp 2008). The presence
of Calliax, today known also from very shallow environments
(Ngoc-Ho 2003) is in accordance with this interpretation.

Fig. 12. Portunoidea gen. et spec. indet. Uncatalogued specimens from Limberg deposited in the Krahuletz-Museum at Eggenburg. Scale
bar equals 5 mm.

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Most decapod occurrences in the Ottnangian deposits thus

represent  rather  shallow-water  components  in  contrast  to
deep-water  assemblages  from  the  Karpatian  deposits  docu-
mented  from  the  Slovak  part  of  the  Vienna  Basin  (Hyžný
&  Schlögl  2011)  and  Slovenian  part  of  the  Styrian  Basin
(Gašparič & Hyžný 2014).

Biogeography: The Ottnangian decapod assemblages of

the Central Paratethys exhibit affinities to the (Proto-)Mediter-
ranean, Boreal and Indo-Pacific regions, suggesting faunal ex-
change  among  these  regions  (Table 1).  Mud  and  ghost
shrimps Jaxea, Calliax and Gourretia are also known from the
modern  Mediterranean  Sea  (Ngoc-Ho  2003).  Jaxea  and Cal-
liax  occurred  in  the  Proto-Mediterranean  Atlantic  Region
(sensu  Harzhauser  et  al.  2002)  already  during  the  Miocene
(Hyžný 2011c) and Oligocene (Hyžný & Gašparič 2014), re-
spectively. In the Central Paratethys, Jaxea kuemeli and Cal-
liax  michelottii  persisted  into  the  latest  Early  Miocene
(Karpatian)  (Hyžný  2011c;  Hyžný  &  Gašparič  2014).  Lio-
carcinus  is  today  widespread  in  the  Mediterranean  (Türkay
2001)  and  is  also  known  from  the  Indo-Pacific  (Manning  &
Holthuis 1981). Its oldest occurrence is Liocarcinus oligocae-
nicus  from  the  Oligocene  of  Paratethys  (Paucă  1929;  Jaroš
1939; Jerzmańska 1967). The first occurrence of Palaeomyra
is from the Oligocene of the (Proto-)Mediterranean (Garassino
et al. 2004). In the Miocene, the genus was already spread and
is known from the Early and Middle Miocene in Central Para-
tethys  (Müller  1984;  this  paper)  and  Late  Miocene  in  the
(Proto-)Mediterranean  (Sismonda  1861;  Garassino  et  al.
2004).  Tanaoa  as  its  supposed  close  relative  (see  above)  is
presently  restricted  to  the  Indo-Pacific  (Galil  2003;  Ng  &
Richer de Forges 2007). Thus, both, Tanaoa and Liocarcinus
may have Tethyan origin. Tethyan ancestry for several deca-
pod  taxa  inhabiting  the  Indo-Pacific  has  already  been  pro-
posed  (for  non-decapod  taxa  see  Harzhauser  et  al.  2007,
2008),  including  the  mud  shrimp  Jaxea  (Hyžný  2011c)  and
the ghost shrimp Glypturus Stimpson, 1866 (Hyžný & Müller
2012).  Although  the  Miocene  fossil  record  of  decapods  in
the  Indo-Pacific  is  moderate  (Noetling  1901;  Böhm  1922;
Van  Straelen  1938;  Morris  &  Collins  1991;  Collins  et  al.
2003; Karasawa et al. 2008b), the Early Miocene appearance
of  Palaeomyra  (? = Tanaoa)  and  Liocarcinus  in  the  circum-
Mediterranean can also imply that the sources of present-day
diversity hotspots trace to the Western Tethys.

Table 1: Occurrences of selected genus-level taxa in space and time. * – denotes the unpublished occurrence (MH pers. obs.)

Oligocene

Early Miocene

Middle Miocene

Late Miocene

Recent

Boreal region

Calliax

Liocarcinus

Central Paratethys

Liocarcinus Jaxea

Calliax
Gourretia
Palaeomyra
Liocarcinus

Jaxea
Calliax
Gourretia*
Palaeomyra
Liocarcinus

(Proto-)Mediterranean

Palaeomyra

Jaxea
Calliax

Jaxea
Palaeomyra
Liocarcinus

Jaxea
Calliax
Gourretia
Liocarcinus

Indo-Pacific

Jaxea
Gourretia
Tanaoa
Liocarcinus

Acknowledgments: We are grateful to Christian Rupp (Geo-
logical  Survey,  Vienna,  Austria)  for  his  help  with  the  strati-
graphy  of  some  of  the  sections  in  Lower  Austria.  Fritz
Steininger  (Krahuletz-Museum  in  Eggenburg,  Austria)  is
thanked for allowing access to crab specimens from Limberg.
Štefan  Józsa  (Comenius  University,  Bratislava,  Slovakia)
helped with processing the figures. Adam Tomašových (Geo-
logical Institute of Slovak Academy of Sciences, Bratislava) is
thanked  for  editorial  comments.  The  critical  remarks  of  two
anonymous reviewers are also acknowledged. The senior au-
thor has been supported by the Austrian Science Fund (FWF;
Lise  Meitner  Program  M  1544-B25).  This  work  was  sup-
ported by the Slovak Research and Development Agency un-
der contracts Nos. APVV-0644-10 and APVV-0436-12.

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