GeolCarp_Vol67_No5_471

GEOLOGICA CARPATHICA

, OCTOBER 2016, 67, 5, 471– 494

doi: 10.1515/geoca-2016-0030

www.geologicacarpathica.com

Diversity and distribution patterns of the Oligocene and

Miocene decapod crustaceans (Crustacea: Malacostraca)

of the Western and Central Paratethys

MATÚŠ HYŽNÝ

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

Department of Geology and Palaeontology, Faculty of Natural Sciences, Comenius University, Ilkovičova 6, 842 15 Bratislava, Slovakia;

hyzny.matus@gmail.com

(Manuscript received June 17, 2015; accepted in revised form September 22, 2016)

Abstract: Decapod associations have been significant components of marine habitats throughout the Cenozoic when

the major diversification of the group occurred. In this respect, the circum-Mediterranean area is of particular interest

due to its complex palaeogeographic history. During the Oligo-Miocene, it was divided in two major areas,

Medi terranean and Paratethys. Decapod crustaceans from the Paratethys Sea have been reported in the literature since

the 19

century, but only recent research advances allow evaluation of the diversity and distribution patterns of

the group. Altogether 176 species-level taxa have been identified from the Oligocene and Miocene of the Western and

Central Paratethys. Using the three-dimensional NMDS analysis, the composition of decapod crustacean faunas of

the Paratethys shows significant differences through time. The Ottnangian and Karpatian decapod associations were

similar to each other both taxonomically and in the mode of preservation, and they differed taxonomically from

the Badenian ones. The Early Badenian assemblages also differed taxonomically from the Late Badenian ones.

The time factor, including speciation, immigration from other provinces and/or (local or global) extinction, can explain

temporal differences among assemblages within the same environment. High decapod diversity during the Badenian

was correlated with the presence of reefal settings. The Badenian was the time with the highest decapod diversity,

which can, however, be a consequence of undersampling of other time slices. Whereas the Ottnangian and Karpatian

decapod assemblages are preserved virtually exclusively in the siliciclastic “Schlier”-type facies that originated in

non-reefal offshore environments, carbonate sedimentation and the presence of reefal environments during

the Badenian in the Central Paratethys promoted thriving of more diverse reef-associated assemblages. In general,

Paratethyan decapods exhibited homogeneous distribution during the Oligo-Miocene among the basins in

the Paratethys. Based on the co-occurrence of certain decapod species, migration between the Paratethys and the North

Sea during the Early Miocene probably occurred via the Rhine Graben. At larger spatial scales, our results suggest that

the circum- Mediterranean marine decapod taxa migrated in an easterly direction during the Oligocene and/or

Miocene, establishing present-day decapod communities in the Indo-West Pacific.

Key words: Decapod crustaceans, Cenozoic, Paratethys, Mediterranean, palaeobiogeography.

Introduction

Decapod associations have been significant components of

marine habitats since the Mesozoic, with ever-increasing

importance throughout Palaeogene and Neogene (Glaessner

1969; Schweitzer 2001; Feldmann & Schweitzer 2006;

Klompmaker et al. 2013) until today (Noël et al. 2014).

Decapods have planktonic larvae (Martin 2014a) and some

of them are active swimmers, so their dispersal and migration

can be very quick (Por 1986). Previously, Müller (1979a)

argued that brachyuran decapods are “among the best zoo-

geographical indicators” because their migration is fast,

many forms can penetrate straits and channels of abnormal

salinity and the duration of their pelagic larval stage is well

defined. Today, the state of knowledge of European Cenozoic

decapods seems to be robust enough to use them for palaeo-

biogeographical studies (Hyžný 2015).

During the Eocene the collision of the African continent

with the European plate initiated the Alpine orogeny, which

resulted in the break-up of the Tethyan Realm into two different

palaeogeographical areas: circum-Mediterranean in the south

and Paratethys in the north (Rögl & Steininger 1983; Rögl

1998). An increase in biogeographical differentiation can be

observed during the Oligocene and Miocene (e.g., Harzhauser

et al. 2002; Harzhauser & Piller 2007). From the Oligocene

onward, the northern domain became a large network of

inland seas intermittently connected to the Mediterranean,

the Atlantic and also to the Indo-Pacific (Rögl & Steininger

1983; Rögl 1998, 1999; Popov et al. 2004; Harzhauser &

Piller 2007; Harzhauser et al. 2007; Kroh 2007; Reuter et al.

2009; Bartol et al. 2014; Fig. 1). For enclosed or semi-

enclosed basins, seaways are very important because they are

the only means of communication with other basins. Varia-

tions in the opening of the seaways were apparently respon-

sible for changes in temperature, salinity and exchange flows

(Popov et al. 2004; Báldi 2006; Moissette et al. 2006; Karami

et al. 2011). Seaways and landbridges discriminate the northern

domain into Western, Central and Eastern Paratethys.

Migration patterns between Paratethys and adjacent areas

of the Western Tethys have been previously investigated

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using foraminifers (Báldi 2006), bryozoans (Moissette et al.

2006), brachiopods (Kocsis et al. 2012), gastropods (Harz-

hauser et al. 2002, 2003, 2007), bivalves (Studencka et al.

1998; Harzhauser et al. 2003, 2007, 2008), and echinoderms

(Kroh & Harzhauser 1999; Harzhauser et al. 2007). To test

these patterns with decapod crustaceans, a database has been

compiled including all previously published Oligocene and

Miocene decapod occurrences and newly gathered data from

unpublished material deposited in institutional collections.

The goals of the present contribution are: 1) to evaluate the

previous research on decapod crustaceans of the Western and

Central Paratethys; 2) to summarize the present knowledge

of decapod associations from the Oligocene and Miocene by

listing all respective species-level taxa and discussing their

occurrences; 3) to discuss factors influencing temporal and

spatial changes of the decapod species richness, using multi-

variate analyses; and 4) to outline distribution patterns of

decapod crustaceans during the Oligo-Miocene.

Study area: Western and Central Paratethys

The studied area comprises basins once belonging to

Western and/or Central Paratethys as indicated in Fig. 2.

The Western Paratethys covered the area of Switzerland,

southern Germany and western Austria. The Central Para-

tethys covered the area from the present-day Austria to

Poland, Ukraine and Romania – it includes the Eastern

Alpine-Carpathian Foreland basins (from Lower Austria to

Moldova) and the area of the Pannonian Basin System itself

formed first during the Early/Middle Miocene.

The geographic separation of the Mediterranean and the

Paratethys seas resulted in a biogeographic differentiation

(Harzhauser et al. 2002; Harzhauser & Piller 2007) and neces-

sitated the establishment of a regional stratigraphic scheme

(Piller et al. 2007). The definition of the regional stages is

based solely on fossil contents (assemblage and abundance

zones). Correlation with Mediterranean/Global chronostrati-

graphy is based mainly on calcareous nannoplankton, plank-

tonic and larger benthic foraminifers (Piller et al. 2007). All

stages are bounded by sea level lowstands coinciding with

order sea level cycles and can be corre lated with the sea

level curve of Haq et al. (1988) and sequence stratigraphic

cycles of Hardenbol et al. (1998). Important summaries of the

Oligocene and Miocene Central Paratethys stratigraphy and

palaeogeography include works by Rögl (1998, 1999), Kováč

et al. (2003, 2007, 2016), Popov et al. (2004, 2006), Piller et

al. (2007) and Harzhauser & Piller (2007).

Material and methods

The synopsis of all occurrences of Oligocene and Miocene

decapod crustaceans is based on two types of sources:

1) specimens and/or taxa examined personally by me and/or

in cooperation with other colleagues; and 2) specimens and/or

Fig. 1. Changing palaeogeography of the circum-Mediterranean

area. Selected time slices include: A — Early Oligocene (ca. 30 Ma);

B — Early Miocene (ca. 17.5 Ma); C — Middle Miocene

(ca. 13 Ma); D — Late Miocene (ca. 8 Ma). Maps modified

from Rögl (1999). CP, EP, WP = Central, Eastern and Western

Paratethys, respectively.

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taxa mentioned in the literature only and still waiting for

re-examination. The material examined personally includes

previously published occurrences as well as still unpublished

collection material. A combination of 10 basins (North

Alpine Foreland Basin = NAFB; Vienna Basin = VB; Styrian

Basin = StB; Hrvatsko Zagorje, Slovenian and Mura basins =

HZB, SlB, MB; Danube Basin = DB; Great Hungarian

Basin = GHB; Novohrad-Nógrad Basin = NNB; Transcar-

pathian Basin = TcB; Transylvanian Basin = TsB; Carpathian

Foreland Basin = CFB) and 6 (sub)stages (Ottnangian, Kar-

patian, Early Badenian, Middle Badenian, Late Badenian,

Early Sarmatian) defines 21 samples. Not all (sub)stages are

represented in every basin. Altogether 176 species-level taxa

have been identified. Their stratigraphic range and geographi-

cal distribution in the Paratethys is indicated in Table 1.

In the analyses of temporal changes in the species compo-

sition (i.e., presence-absence), pre-Karpatian, Sarmatian

(two samples) and samples with less than three species were

removed from the compositional matrix (modified Table 1).

The three-dimensional non-metric multidimensional scaling

(NMDS) and cluster analysis have been performed using

freeware PAST (Hammer et al., 2001). In addition, decapod

associations from reefal and non-reefal settings were

analysed separately. Reefal taxa are defined here as taxa

occurring exclusively in reefal settings. The Jaccard index

was used to quantify dissimilarity among analysed associa-

tions. Because most decapod occurrences come from

non-quantitative samples, statistical comparison of species

abundances was not possible.

Major collections of the Miocene and Oligocene fossil

decapod crustaceans, which formed the basis of the database,

are housed in several European institutions. All of them

except the Polish Academy of Sciences Museum of the Earth

in Warsaw were visited and studied first-hand. Abbreviations

of the repository institutions: GBA — Geological Survey in

Vienna, Austria; HNHM — Hungarian Natural History

Museum in Budapest, Hungary; KGP-MH — Department of

Geology and Palaeontology, Comenius University in

Bratislava, Slovakia; MFGI — Hungarian Geological and

Geophysical Survey in Budapest, Hungary; MTM — Mátra

Museum in Gyöngyös, Hungary; MZ — Polish Academy of

Sciences Museum of the Earth in Warsaw, Poland; NHMW

— Geological-palaeontological Department, Natural History

Museum in Vienna, Austria; RGA/SMNH — “Rok Gašparič

palaeontological collection”, Slovenian Museum of Natural

History in Ljubljana, Slovenia; SNM-Z — Natural History

Museum of the Slovak National Museum in Bratislava, Slovakia;

UMJGP — Universalmuseum Joanneum in Graz, Austria. In the

following text, regional Paratethyan stages are largely used.

For correlation with the Mediterranean scale, see Fig. 3.

Results

Decapod associations basin by basin

Decapod crustaceans from the area once covered by the

Paratethys Sea have been reported in the literature since the

century. The first papers were dedicated to sporadic fin-

dings of mainly Middle Miocene taxa. Among the first scholars

studying Paratethyan decapods were August Reuss (*1811–

†1873), Alexander Bittner (*1850 –†1902) and Paul Brocchi

(*1838 –†1898). Subsequently, in the first half of the

century, Imre (Emmerich) Lőrenthey (*1867 –†1917; see

Fig. 2. Neogene basins of the Western and Central Paratethys analysed in this study. White colour represents the extent of exposed strata of

respective basins. CFB = Carpathian Foreland Basin, DB = Danube Basin, GHB = Great Hungarian Basin, HZB = Hrvatsko-Zagorje Basin,

MB = Mura Basin, NAFB = North Alpine Foreland Basin, NNB = Novohrad-Nógrad Basin, StB = Styrian Basin, TcB = Transcarpathian

Basin, TsB = Transylvanian Basin, VB = Vienna Basin.

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Table 1: List of the Western and Central Paratethys decapod species with occurrence within the studied basins (codes for basins used as in

Fig. 1 and the text). Freshwater taxa are not included. Asterisk (*) indicates the species exhibiting extra-Paratethyan distribution.

Excla mation (!) indicates the species living until today. Abbreviations (taxonomy) used: AL = Albuneidae, AN = Anomura, AX = Axiidea,

BR = Brachyura, CA = Callianassidae, CAL = Calappoidea, CAN = Cancroidea, CAR = Carpilioidea, CT = Ctenochelidae, DOR = Dorripoidea,

DRO = Dromioidea, GA = Galatheoidea, GE = Gebiidea, GON = Goneplacoidea, GRA = Grapsoidea, LA = Laomediidae, LEU = Leucosioidea,

MAJ = Majoidea, PAL = Palicoidea, PAR = Parthenopoidea, PG = Paguroidea, PIL = Pilumnoidea, POR = Portunoidea, RAN = Raninoidea,

RET = Retroplumoidea, TH = Thomassiniidae, UP = Upogebiidae, XAN = Xanthoidea.

Taxon (species)

Kiscellian / Rupelian

Egerian / Chattian

Eggenburgian

Ottnangian / Helvetian

Karpatian

E. Badenian

M. Badenian

L. Badenian

E. Sarmatian

GE/UP

Upogebia scabra Müller, 1974b

GHB

GE/LA

Jaxea kuemeli Bachmayer, 1954

NAFB NNB,

StB

NNB,

StB

AX/CA

* “Callianassa” almerai Müller, 1993

StB

AX/CA “Callianassa” ferox Bittner, 1893

TsB

AX/CA “Callianassa” jahringensis Glaessner, 1928

StB

AX/CA “Callianassa” kerepesiensis Müller, 1976

NNB

GHB

AX/CA

* “Callianassa” cf. kerepesiensis Müller, 1976

CFB,

NNB

GHB

AX/CA “Callianassa” norica Glaessner, 1928

StB

AX/CA “Callianassa” oroszyi (Bachmayer, 1954)

AX/CA “Callianassa” rapax Bittner, 1893

TsB

AX/CA “Callianassa” roztoczensis Müller, 1996

CFB,

GHB

AX/CA “Callianassa” simplex Bittner, 1893

TsB

AX/CA “Callianassa” velox Bittner, 1893

TsB

AX/CA “Callianassa” vorax Bittner, 1893

TsB

AX/CA

*Calliax michelottii (A. Milne Edwards, 1860)

NAFB

NAFB StB

NNB,

StB

AX/CA

Eucalliax pseudorakosensis (Lőrenthey in Lőrenthey &

Beurlen, 1929)

NNB TsB

GHB,

AX/CA

*Calliaxina chalmasii (Brocchi, 1883)

GHB VB

AX/CA

Callichirus bertalani Hyžný & Müller, 2010a

AX/CA

Neocallichirus brocchii (Lőrenthey, 1897)

GHB VB

AX/CA

Balsscallichirus florianus (Glaessner, 1928)

GHB

StB,

StB

AX/CA

*Balsscallichirus sismondai (A. Milne Edwards, 1860)

StB

AX/CA

*Glypturus munieri (Brocchi, 1883)

DB,

StB

DB,

GHB,

AX/CA

Lepidophthalmus crateriferus (Lőrenthey in Lőrenthey &

Beurlen, 1929)

GHB

AX/CA

Lepidophthalmus paratethyensis Gašparič & Hyžný, 2014

StB

AX/CT

*Ctenocheles rupeliensis (Beurlen, 1939)

GHB

AX/CT

Callianopsis marianae Hyžný & Schlögl, 2011

AX/TH

Crosniera schweitzerae Hyžný & Schlögl, 2011

AN/PG

Anapagurus carinatus Harvey, 1998

GHB

AN/PG

Anapagurus miocenicus Müller, 1978

StB

GHB

AN/PG

Ciliopagurus substriatiformis (Lőrenthey in Lőrenthey &

Beurlen, 1929)

GHB,

AN/PG !Dardanus arrosor (Herbst, 1796)

AN/PG

*Dardanus hungaricus (Lőrenthey in Lőrenthey & Beurlen,

1929)

CFB,

DB,

NNB,

StB

GHB

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Table 1 (continuation):

Taxon (species)

Kiscellian / Rupelian

Egerian / Chattian

Eggenburgian

Ottnangian / Helvetian

Karpatian

E. Badenian

M. Badenian

L. Badenian

E. Sarmatian

AN/PG !Diogenes cf. pugilator (Roux, 1828)

GHB

AN/PG

Diogenes matrensis Müller, 1984a

NNB

AN/PG

Paguristes cserhatensis Müller, 1984a

NNB

AN/PG

Pagurus concavus Müller, 1978

GHB

AN/PG

Pagurus rakosensis Müller, 1978

CFB,

GHB

AN/PG

Pagurus retznensis Collins, 2014

StB

AN/PG

Pagurus tuberculosus Harvey, 1998

GHB

AN/PG

Pagurus turcus Müller, 1984a

AN/PG

*Petrochirus priscus (Brocchi, 1883)

StB

TsB

GHB,

AN/PG

Pylopagurus corallinus Müller, 1996

CFB,

NNB

AN/PG

Pylopagurus leganyii Müller, 1984a

NNB

AN/GA

Agononida cerovensis Hyžný & Schlögl, 2011

AN/GA

Galathea weinfurteri Bachmayer, 1950

CFB,

GHB

CFB,

AN/GA

Munidopsis lieskovensis Hyžný & Schlögl, 2011

AN/GA

Munidopsis palmuelleri Hyžný, Gašparič, Robins & Schlögl,

2014

StB

AN/GA

Petrolisthes haydni Müller, 1984a

StB

CFB

AN/GA

Petrolisthes magnus Müller, 1984a

CFB,

GHB

AN/GA

*Pisidia kokayi (Müller, 1974a)

NNB VB

GHB

AN/GA

*Pisidia viai Müller, 1984b

DB,

StB

AN/GA

Pisidia aff. viai Müller, 1984b

CFB

AN/GA

Pisidia? subnodosa Collins, 2014

StB

AN/AL

Albunea asymmetrica Müller, 1978

GHB

55 BR/DRO

Dromia evae Collins, 2014

StB

56 BR/DRO

*Dromia neogenica Müller, 1978

CFB,

GHB

57 BR/DRO

Dynomene emiliae Müller, 1979b

CFB,

CFB

58 BR/DRO

Kerepesia viai Müller, 1976

StB

GHB

59 BR/DRO

Kromtitis koberi (Bachmayer & Tollmann, 1953)

CFB,

DB,

StB

60 BR/DRO

Lucanthonisia eotvoesi (Müller, 1975a)

CFB,

GHB

61 BR/DOR

Dorippe fankhauseri Studer, 1892

NAFB

62 BR/DOR

Dorippe margaretha Lőrenthey in Lőrenthey & Beurlen, 1929

GHB

63 BR/DOR

Dorippe ornatissima Müller, 2006

GHB

64 BR/DOR

Neodorippe carpathica (Förster, 1979a)

CFB

65 BR/DOR

Ethusa octospinosa Müller, 2006

GHB

66 BR/RAN

Lyreidus hungaricus Beurlen, 1939

GHB

67 BR/RAN

Ranidina rosaliae Bittner, 1893

CFB,

68 BR/CAL

Calappa heberti Brocchi, 1883

GHB,

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Table 1 (continuation):

Taxon (species)

Kiscellian / Rupelian

Egerian / Chattian

Eggenburgian

Ottnangian / Helvetian

Karpatian

E. Badenian

M. Badenian

L. Badenian

E. Sarmatian

69 BR/CAL

*Calappa praelata Lőrenthey in Lőrenthey & Beurlen, 1929

CFB,

DB,

NNB,

StB,

CFB,

70 BR/CAL

Calappilia matzkei (Bachmayer, 1962)

71 BR/CAL

Calappilia tridentata (Beurlen, 1939)

GHB

72 BR/CAL

Mursia harnicari Hyžný & Schlögl, 2011

73 BR/CAL

*Mursia lienharti (Bachmayer. 1962)

GHB,

74 BR/CAL

Osachila tiechei (Studer, 1898)

NAFB

75 BR/CAL

Szaboa inermis (Brocchi, 1883)

GHB VB

76 BR/CAN

Cancer illyricus Bittner, 1883

HZB

77 BR/CAN

Cancer styriacus Bittner, 1884

StB

CFB,

78 BR/CAN

Corystites latifrons (Lőrenthey in Lőrenthey & Beurlen, 1929)

GHB

79 BR/CAN

Glebocarcinus helveticus Fraaije, Menkveld-Gfeller,

van Bakel & Jagt, 2010

NAFB

80 BR/CAN

*Lobocarcinus sismondai von Meyer, 1843

81 BR/CAN

Lobocarcinus aff. sismondai von Meyer, 1843

StB

82 BR/CAN

Microdium nodulosum Reuss, 1867

CFB

83 BR/CAN

Miocyclus bulgaricus Müller, 1979b

CFB

84 BR/CAN

*Tasadia carniolica (Bittner, 1884)

NNB,

GHB,

SlB,

85 BR/CAR

*Eocarpilius antiquus (Glaessner, 1928)

CFB VB

86 BR/GON

Coeloma macoveii Lăzărescu, 1959

TsB

87 BR/GON

Coeloma egerense Lőrenthey in Lőrenthey & Beurlen, 1929

GHB

88 BR/GON

*Goneplax gulderi Bachmayer, 1953c

StB

CFB

89 BR/GON

Goneplax? carnuntinus (Bachmayer, 1953b)

90 BR/GON

Mioplax socialis Bittner, 1884

CFB,

HZB,

StB,

TcB

91 BR/GON

Neopilumnoplax pohorjensis Gašparič & Hyžný, 2014

StB

92 BR/GON

Styrioplax exiguus (Glaessner, 1928)

NNB,

StB

93 BR/LEU

Andorina elegans Lőrenthey, 1901

GHB

94 BR/LEU

Ebalia hungarica Müller, 1974a

GHB

95 BR/LEU

Ebalia oersi Müller, 1978

GHB

96 BR/LEU

Ebalia vanstraeleni Bachmayer, 1953b

97 BR/LEU

Myra emarginata Glaessner, 1928

StB

98 BR/LEU

Palaeomyra globulosa (Müller, 1975a)

GHB

99 BR/LEU

Palaoemyra strouhali (Bachmayer, 1953a)

NAFB

100 BR/MAJ

Achaeus magnus Müller, 1978

GHB

101 BR/MAJ

Hyas meridionalis Glaessner, 1928

StB

102 BR/MAJ

*Maja biaensis Lőrenthey in Lőrenthey & Beurlen, 1929

CFB,

GHB

103 BR/MAJ

Micippa hungarica (Lőrenthey in Lőrenthey & Beurlen, 1929)

GHB,

104 BR/MAJ

Pisa oroszyi (Bachmayer, 1953b)

CFB,

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Table 1 (continuation):

Taxon (species)

Kiscellian / Rupelian

Egerian / Chattian

Eggenburgian

Ottnangian / Helvetian

Karpatian

E. Badenian

M. Badenian

L. Badenian

E. Sarmatian

105 BR/MAJ

*Schizophrys visegradensis Müller, 1984a

106 BR/PAL

Crossotonotus diosdensis Müller, 1984a

GHB

107 BR/PAL

Palicus hungaricus Müller, 2006

GHB

108 BR/PAR

Parthenope szaboi Müller, 1974b

GHB,

109 BR/PAR

Parthenope tetenyensis Müller, 1984a

GHB

110 BR/PIL

Actumnus telegdii (Müller, 1974b)

CFB,

GHB

111 BR/PIL

Glabropilumnus fossatus Müller, 1996

CFB,

112 BR/PIL

Glabropilumnus nitidus Collins, 2014

StB

113 BR/PIL

Pilumnopeus paratethyensis Müller, 1984a

GHB

114 BR/PIL

Pilumnopeus tetenyensis Müller, 1984a

GHB

115 BR/PIL

Pilumnus mediterraneus (Lőrenthey, 1897)

CFB,

NNB

CFB,

GHB

116 BR/PIL

Pilumnus aff. mediterraneus (Lőrenthey, 1897)

StB

117 BR/POR

Bathynectes muelleri Ossó & Stalennuy, 2011

CFB

118 BR/POR !Carupa cf. tenuipes Dana, 1852

119 BR/POR

Chaceon heimertingensis (Bachmayer & Wagner, 1957)

NAFB

120 BR/POR

Charybdis fragilis (Müller, 1978)

StB

GHB

121 BR/POR

Charybdis mathiasi Müller, 1984a

GHB

122 BR/POR

*Geryon cf. latifrons Van Straelen, 1937

GHB

123 BR/POR

Liocarcinus kuehni (Bachmayer, 1953b)

GHB,

124 BR/POR

Liocarcinus oligocaenicus (Paucă, 1929)

CF,

TsB

GHB

125 BR/POR

Liocarcinus oroszyi (Bachmayer, 1953b)

126 BR/POR

Liocarcinus ottnangensis (Bachmayer, 1953a)

NAFB

127 BR/POR

Liocarcinus praearcuatus Müller, 1996

CFB,

GHB

128 BR/POR

Liocarcinus rakosensis (Lőrenthey in Lőrenthey & Beurlen,

1929)

NNB TsB,

CFB,

GHB

129 BR/POR

Liocarcinus aff. rakosensis (Lőrenthey in Lőrenthey &

Beurlen, 1929)

StB

130 BR/POR

Lissocarcinus szoeraenyiae (Müller, 1974b)

GHB

131 BR/POR

Macropipus gruneri Bachmayer & Rutsch, 1962

NAFB

132 BR/POR

Miopipus pygmeus (Brocchi, 1883)

GHB

133 BR/POR

Mioxaiva psammophila Müller, 1978

GHB

134 BR/POR

Necronectes schafferi Glaessner, 1928

StB,

GHB

135 BR/POR

Pirimela loerentheyi (Müller, 1974a)

GHB

136 BR/POR

Portumnus tricarinatus (Lőrenthey in Lőrenthey & Beurlen,

1929)

GHB

137 BR/POR

Portunus kisslingi Studer, 1892

NAFB

138 BR/POR

Portunus krambergeri Bittner, 1893

HZB

139 BR/POR

Portunus miocaenicus Müller, 1984a

GHB

140 BR/POR

*Portunus monspeliensis (A. Milne Edwards, 1860)

NNB,

StB

GHB

141 BR/POR

Portunus muelleri Collins, 2014

StB

142 BR/POR

Portunus neogenicus Müller, 1978

GHB

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Table 1 (continuation):

Taxon (species)

Kiscellian / Rupelian

Egerian / Chattian

Eggenburgian

Ottnangian / Helvetian

Karpatian

E. Badenian

M. Badenian

L. Badenian

E. Sarmatian

143 BR/POR

Portunus radobojanus (Bittner, 1884)

HZB

144 BR/POR

Portunus stenaspis (Bittner, 1884)

HZB

145 BR/POR

*Portunus viai Secretan in Philippe & Secretan, 1971

GHB

146 BR/POR

Rakosia carupoides Müller, 1984a

CFB,

GHB

147 BR/POR

Rakosia rectifrons Müller, 1996

CFB

148 BR/POR

Scylla molassica Studer, 1898

NAFB

149 BR/POR

*Scylla michelini A. Milne Edwards, 1860

StB

150 BR/POR

Trachypirimela grippi (Müller, 1974b)

GHB

151 BR/POR

Xaiva bachmayeri Müller, 1984a

GHB

152 BR/RET

*Loerenthopluma lata Beschin, Busulini, De Angeli &

Tessier, 1996

GHB

153 BR/RET

Retropluma slovenica Gašparič & Hyžný, 2014

StB

NNB

154 BR/XAN

*Actaea turcocampestris Müller, 1984a

DB,

StB,

155 BR/XAN

*Daira speciosa (Reuss, 1871)

DB,

StB,

CFB,

GHB

156 BR/XAN

Eomaldivia friebei Collins, 2014

StB

157 BR/XAN

*Etisus evamuellerae Hyžný, Van Bakel & Guinot, 2014

158 BR/XAN

Glyphithyreus sulcatus (Beurlen, 1939)

GHB

159 BR/XAN

Haydnella steiningeri Müller, 1984a

CFB,

DB,

StB

160 BR/XAN

Chlorodiella juglans Müller, 1984a

StB

CFB,

GHB

161 BR/XAN

Chlorodiella loczyi Müller, 1984a

StB

GHB

162 BR/XAN

Chlorodiella mediterranea (Lőrenthey in Lőrenthey &

Beurlen, 1929)

CFB,

DB,

StB

GHB

163 BR/XAN

*Chlorodiella tetenyensis Müller, 1984a

StB

CFB,

GHB

164 BR/XAN

Jonesius planus (Müller, 1996)

CFB VB

165 BR/XAN

Panopeus wronai Müller, 1984a

StB

CFB

166 BR/XAN

Pilodius vulgaris (Glaessner, 1928)

GHB

167 BR/XAN

Trapezia glaessneri Müller, 1975b

168 BR/XAN

*Xantho moldavicus (Yanakevich, 1977)

StB

CFB,

GHB,

169 BR/GRA

Asthenognathus rakosensis Müller, 2006

GHB

170 BR/GRA

Brachynotus februarius Müller, 1974a

GHB

171 BR/GRA

Metopograpsus badenis Müller, 2006

GHB

172 BR/GRA

Metopograpsus traxleri Müller, 1998

173 BR/GRA

Pachygrapsus hungaricus Müller, 1974a

CFB,

GHB

174 BR/OCY

*Macrophthalmus aquensis A. Milne Edwards & Brocchi,

1879

CFB,

StB

175 BR/OCY

Paracleistostoma miocaenica Müller, 1998

176 BR/OCY

Tritodynamia miocaenica Müller, 2006

GHB

Total number of species

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Müller 1987), Karl Beurlen (*1901 –†1985) and Martin F.

Glaessner (*1906 –†1989; see Küpper 1991) contributed to

the study of fossil decapod crustaceans with several mile-

stone works (Lőrenthey & Beurlen 1929; Glaessner 1929,

1969). In the second half of the 20

century Friedrich

Bachmayer (*1913 –†1989; see Kollmann & Zapfe 1979),

Reinhard Förster (*1935 –†1987; see Herm 1987) and Pál

Müller (*1935 –†2015; see Hyžný et al. 2014a, 2015a) pub-

lished numerous papers focused (mostly) on Neogene deca-

pods of the Western and Central Paratethys.

All decapod species (excluding freshwater taxa) reported

from the respective area are listed in Table 1. Proportions of

decapod species per stage and basin are shown in Figs. 3 and

4, respectively.

North Alpine Foreland Basin: From the Oligocene

(Chattian) of Bavaria, Germany, Bachmayer & Wagner

(1957) recorded a single occurrence of Chaceon heimertin

gensis (Bachmayer & Wagner, 1957). From the Oligo-

Miocene (Chattian-Aquitanian) of Bavaria, Barthelt (1989)

mentioned claws attributed to Callianassa sp. More decapod

associations are known from the Molasse Zone of Switzer-

land; Fröhlicher (1951) reported on portunid crabs from

the Oligocene strata, whereas Studer (1892, 1898) and

Bachmayer & Rutsch (1962) described several taxa from the

Miocene “Meeresmolasse”. Fraaije et al. (2010a) briefly

reviewed all Lower Miocene decapod crustaceans from the

type area of the Helvetian stage in the Bern area (Switzer-

land), including one new species, Glebocar

cinus helveticus

Fraaije, Menkveld-Gfeller, van Bakel & Jagt, 2010.

Bachmayer (1953a, 1982) reported several brachyuran

species from the Ottnangian strata of Ottnang and Limberg,

respectively. Hyžný (2011a) reported Jaxea kuemeli

Bachmayer, 1954 from the same site. All decapod taxa

known from the Ottnangian strata of Austria and Germany

have been revised by Hyžný et al. (2015b).

Vienna Basin and its satellite basins: The Miocene

deposits of the Vienna Basin and its satellite basins have

yielded many decapod crustacean remains; altogether

65 species were identified in this basin (Fig. 4). A decapod

fauna from the Karpatian of the Korneuburg Basin was

described by Glaessner (1924, 1928), Müller (1998) and

Hyžný (2016). Hyžný & Schlögl (2011) described roughly

coeval association from the northern Vienna Basin. Most

decapod remains of the Vienna Basin are of Badenian age

and were described by many authors (Reuss 1859; Von Reuss

1871; Bittner 1877; Toula 1904; Glaessner 1928; Bachmayer

1950, 1953b, c, 1954, 1962; Bachmayer & Küpper 1952;

Bachmayer & Tollmann 1953; Müller 1984a). The localities

were summarized by Müller (1984a).

Styrian Basin: Decapod crustaceans from the Miocene

deposits of the Austrian part of the Styrian Basin were

reported by Glaessner (1928), Schouppe (1949), Flügel

(1986) and Friebe (1987, 1990). Badenian reef-associated

Fig. 3. Decapod crustacean species diversity of the Central and

Western Paratethys per stage. Oligo-Miocene stages correlated to

regional chronostratigraphy of the Central Paratethys are adapted

from Piller et al. (2007). Absolute ages are based on the Interna-

tional Chronostratigraphic Chart (www.stratigraphy.org).

Fig. 4. Proportion of the studied decapod taxa per basin. Basin

abbreviations used as in Fig. 1.

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decapod fauna of the quarry in Retznei (Reuter & Piller 2011;

Reuter et al. 2012) has been described by Collins (2014);

Hyžný (2011b) reported from the same locality ghost shrimps

preserved in situ within the burrow structures. Several ghost

shrimps and few brachyurans were described from the Kar-

patian sediments cropping out at localities close to Austria/

Slovenia border (Glaessner 1928). From the Slovenian part

of the Styrian Basin Miocene decapod faunas were reported

by Glaessner (1928), Mikuž (2003a), Gašparič & Hyžný

(2014) and Gašparič & Halásová (2015).

Hrvatsko Zagorje, Slovenian and Mura basins: Spo-

radic finds of the Oligocene and Miocene decapods from the

Slovenian Basin were reported by Bittner (1884), Mikuž

(2003b, 2010), Mikuž & Pavšič (2003), and Križnar (2009).

Decapod faunas from the Karpatian of the Mura Basin were

described by Glaessner (1928). Miocene decapods from the

Hrvatsko Zagorje Basin (Croatia) were reported by Bittner

(1884, 1893); Bittner (1893) described Achelous krambergeri

(now treated as Portunus) from the Egerian of Varaždin and

Bittner (1884) described a small decapod fauna from the

Badenian of Radoboj.

Danube Basin: Müller (1984a) described several localities

in the Hungarian part of the Danube Basin (Börzsöny near

Visegrád) with Badenian decapod remains. Hyžný & Müller

(2010a) described additional ghost shrimps from the area

close to Lake Balaton. Little is known about the decapods

from the Slovak part of the basin; Špinar et al. (1965) illus-

trated few specimens of Calliax michelottii (A. Milne

Edwards, 1860) (see also Hyžný & Gašparič 2014) and

Goneplax sp. from boreholes near Želiezovce village.

Great Hungarian Basin: Oligo-Miocene strata (and its

“predecessor”, namely the Hungarian Palaeogene Basin)

have yielded 86 species of decapods (Fig. 4). Concerning

decapod crustaceans, this basin is the most sampled area of

the Central Paratethys. A deep-water decapod crustacean

fauna was described by Beurlen (1939) and partly rede-

scribed by Hyžný & Dulai (2014) in the Lower Oligocene

Kiscell Clay Formation. A rare find of Loerenthopluma lata

Beschin, Busulini, De Angeli & Tessier, 1996 from the Oli-

gocene Mány Formation was reported by Hyžný & Müller

(2010b).

From the Upper Oligocene (Egerian) of Eger a new

crab species Coeloma egerense was described by Lőrenthey

in Lőrenthey & Beurlen (1929). The Middle Miocene (Bade-

nian) decapod faunas were extensively documented by

Brocchi (1883), Bittner (1893), Lőrenthey (1897, 1898 a, b, c,

1904 a, b, 1911, 1913), Lőrenthey & Beurlen (1929) and

Müller (1974 a, b, 1975 a, b, 1976, 1978, 1984 a, 2006). Loca-

lities in the Budapest area are in this respect the classical

ones (for details see Müller 1984a).

Novohrad-Nógrad Basin: Few decapod remains of Jaxea

kuemeli and Styrioplax exiguus are known from the Karpa-

tian of the Hungarian part of the basin (Álsoszuha 1 borehole;

see Hyžný 2011a; Hyžný & Schlögl 2011). Müller (1984 a)

described several sites with Badenian decapod remains in the

Hungarian part of the basin; a decapod fauna from the Bade-

nian of the Slovak part of the basin (locality Plášťovce) was

described by Hyžný et al. (2015c).

Transcarpathian Basin: Hyžný & Ledvák (2014) and

Hyžný et al. (2016) reported on the occurrence of Mioplax

socialis Bittner, 1884 from the Sarmatian of the Slovak part

of the basin. No other decapod occurrences from TcB are

known to date.

Transylvanian Basin: Bittner (1893) described small

decapod faunas from the Oligocene Mera Beds and the

Miocene of the vicinity of Cluj (Klausenberg). Lăzărescu

(1959) described Coeloma macoveii Lăzărescu, 1959 from

the Chattian (Early Egerian) of Romania, although

Schweitzer et al. (2009: 11) stated the Miocene age of

the species.

Carpathian Foreland Basin: Oligocene crabs were

described by Paucă (1929), Jaroš (1939) and Jerzmańska

(1967) from Romanian, Moravian and Polish parts of the

Silesian-Krosno Belt, respectively. This belt belonged to the

Silesian-Krosno domain sensu Kováč et al. (2016), a prede-

cessor of the Carpathian Foredeep. Reuss (1867), Förster

(1979a, b), Müller (1984a, 1996) and Górka (2002) described

Badenian decapod associations from several localities

situa ted in the Polish part of the Carpathian Foreland Basin.

A Late Badenian reef-associated decapod fauna of

the Medobory Hills (Ukraine) has been reported by

Radwański et al. (2006), Ossó & Stalennuy (2011) and Górka

et al. (2012).

Freshwater decapods: Fossil freshwater decapods are in

general rare. Only few occurrences are known from the Oligo-

Miocene strata of the studied area. Houša (1956) and

Zázvorka (1956) reported on freshwater shrimps from Czech

Republic and Klaus & Gross (2010) provided a compre-

hensive synopsis of the fossil freshwater crabs of Europe,

including those from the North Alpine Foreland Basin and

Pannonian Basin System.

Analyses

Species richness per stage: The Badenian was the time of

the highest decapod diversity: 90, 73, and 46 species were

recorded from the Late Badenian (B3), Early Badenian (B1)

and Middle Badenian (B2), respectively. Large percentages

of the Badenian taxa, namely 47.9 % (B1), 60.9 % (B2) and

27.8 %, (B3), are known from reefal settings only (Fig. 3).

The smallest diversity was recorded from the Eggenburgian

and Egerian, with only 1 and 7 recorded decapod species,

respectively. The species richness of the Ottnangian and

Sarmatian is similarly low, with ten and nine species,

respectively.

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Species richness per basin: The Great Hungarian Basin,

Vienna Basin, Styrian Basin, and Carpathian Foreland Basin

were relatively well-sampled with 86, 65, 49, and 44 species,

respectively (Fig. 4). Relatively high diversity was observed

always in the basins with the presence of reefal settings. Low

species richness (<12) was recorded in the basins with no

reefal settings. From the Transcarpathian Basin, a single

species was recorded.

NMDS and cluster analyses: Decapod associations of

five time slices were analysed (NMDS, Jaccard index):

Ottnangian (O), Karpatian (K), Early Badenian (B1), Middle

Badenian (B2) and Late Badenian (B3). Ottnangian and Kar-

patian assemblages do not overlap with the Badenian ones.

Early Badenian assemblages were always separated from the

Late Badenian ones without any overlap (Fig. 5). Decapod

taxa restricted to reefal settings analysed separately (NMDS,

Jaccard index) show distinct pattern with associations from

all three substages separated from each other (Fig. 6A).

Cluster analysis (Jaccard index), however, did not resolve the

associations of respective substages so unambiguosly

(Fig. 6B). Interestingly, the Middle Badenian association of

the Vienna Basin (B2-VB) was clustered together with the

Early Badenian association of the Danube Basin (B1-DB),

and the Early Badenian association of the Styrian Basin

(B1-StB) was clustered together with the Late Badenian

associations (B3-GHB, B3-CFB).

The NMDS analysis (Jaccard index) of non-reefal taxa

resolved the Ottnangian and Karpatian associations as

distinct from all Badenian ones (Fig. 7). All Early Badenian

associations were clearly separated from the Late Badenian

ones.

Discussion

Collection bias

A sampling-standardized comparison of decapod species

richness across the basins and/or time intervals is not possible

because some basins and stages are under-represented in the

analysed matrix due to uneven sampling effort, outcrop

availability, facies distribution and preservation potential of

decapod crustaceans. For example, there are only 12, 7 and 1

decapod species recorded from the Kiscellian, Egerian and

Eggenburgian, respectively (Fig. 3). The time span of these

three stages is longer than the time span of the Ottnangian,

Karpatian, Badenian and Sarmatian taken together, and the

smaller diversity of decapod species recorded in these three

stages is probably related to much smaller collection efforts.

More than 60 decapod species were recorded both from the

Great Hungarian Basin and the Vienna Basin. Although

reefal settings in these areas yielded numerous decapod taxa

(Fig. 4), there is at least one more factor contributing to the

observed pattern of species richness. Two cities, Budapest

and Vienna, with a long tradition in fossil decapod research

(see above), are located in the respective basins, presumably

owing to the over-sampling in comparison with other areas.

Moreover, the presence of numerous outcrops with the Bade-

nian strata in the vicinity of Budapest and Vienna contributed

to preferential focus of the scholars on these strata. Thus, the

high decapod species diversity in the Late Badenian of the

Great Hungarian Basin is partially influenced by the scien-

tific contributions of Imre Lőrenthey and Pál Müller, whereas

high diversity of decapod crustaceans in the Vienna Basin

owes much to the efforts of Friedrich Bachmayer (see above).

Fig. 5. NMDS plots of decapod associations of the Western and Central Paratethys using the Jaccard index (stress: 0.195). Labelled points

represent all decapod species per stage and basin. For comments see text. Age abbreviations: O = Ottnangian, K = Karpatian, B1 = Early

Badenian, B2 = Middle Badenian, B3 = Late Badenian. Basin abbreviations used as in Fig. 1.

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Additionally, as stated by Müller (1979a), decapod fossils

“need a careful collecting and preparing process and they are

not very abundant”.

Preservation aspects

Preservation potential of decapod crustaceans is relatively

low compared with molluscs, a major group often used for

investigation of migration patterns (Studencka et al. 1998;

Harzhauser et al. 2002, 2003, 2007, 2008). Bodies of deca-

pod crustaceans are the subject of quick decomposition and

the fragile exoskeleton is often fragmented before burial

takes place (Schäfer 1951; Plotnick 1986; Plotnick et al.

1988; Stempien 2005; Mutel et al. 2008). Ideal conditions

for preservation of complete decapod bodies include a calm

depositional setting coupled with a quick burial without sub-

sequent physical disturbance or biotic reworking (Müller et

al. 2000). Dysoxic and anoxic conditions in the Oligocene

menilites yielded near-complete body fossils of decapods

(Paucă 1929; Jerzmańska 1967; Jaroš 1939; Bieńkowska-

Wasiluk 2010). However, they are flattened, and superim-

posed body parts are difficult to interpret. Moreover, these

occurrences have so far received virtually no or limited taxo-

nomic or taphonomic attention (Glaessner 1965). Thus, it is

not surprising that the observed species richness of the

Oligocene Paratethyan crabs is rather low (Fig. 3).

The Miocene fine-grained siliciclastic facies (“Schlier”-

type and “Tegel”-type facies) often yielded near-complete

decapod specimens. This is the case of the Ottnangian (Bach-

mayer 1982; Hyžný et al. 2016), Karpatian (Hyžný & Schlögl

2011; Gašparič & Hyžný 2015) and some Badenian strata

(e.g., Bittner 1884). Fine-grained volcanoclastics may

promote preservation of complete decapod body fossils as is

the case of the locality Plášťovce (NNB) reported by Hyžný

et al. (2015c). Coarser-grained facies (silici- or volcanoclas-

tic) yielded mostly isolated cheliped fingers and fragmentary

carapaces, which, however, can be successfully taxono-

mically evaluated if thorough comparison with extant taxa is

made (e.g., Müller 1984a; Hyžný & Klompmaker 2015).

Fig. 6. NMDS plot (A) and dendrogram of the cluster analysis (B) of reef-associated decapod associations of the Western and Central

Paratethys using the Jaccard index (stress: 0.113). For comments see text. Age abbreviations: B1 = Early Badenian, B2 = Middle Badenian,

B3 = Late Badenian. Basin abbreviations used as in Fig. 1.

Fig. 7. NMDS plot of non-reefal decapod associations of the West-

ern and Central Paratethys using the Jaccard index (stress: 0.267).

For comments see text. Age abbreviations: O = Ottnangian, K = Kar-

patian, B1 = Early Badenian, B2 = Middle Badenian, B3 = Late Bade-

nian. Basin abbreviations used as in Fig. 1.

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Reefs have an uneven surface and contain a lot of cavities

and micro-lagoons, protected from currents and washout.

Decapod crustaceans are usually preserved in such places,

preferentially in the cavities or pockets sensu Müller (2004).

Thus, even fragile decapod remains tend to be preserved and

have potential to be taxonomically evaluated. It can be con-

cluded that relatively high species diversity observed across

stages (Fig. 3) and basins (Fig. 4) is partly dependent on

the occurrence of reefal settings.

Temporal patterns in the regional species richness

Oligocene: At the beginning of the Oligocene, the Central

Paratethys was separated from the Mediterranean (Fig. 1A).

Individual basins in the Central Paratethys gained a character

of a semi-closed sea (Kováč et al. 2016). They were charac-

terized by primarily estuarine water circulation pattern with

recurrent episodes of stagnation (Popov et al. 2004). As

a consequence, accumulations of dysoxic to anoxic sedi-

ments were predominant during the Oligocene and Early

Miocene in the entire Paratethys (Popov et al. 2004; Kováč et

al. 2016). In the deeper part of the Palaeogene Hungarian

Basin, laminated black shales were deposited in the anoxic

environment represented by the Tard Clay (Báldi 1984) and

Schöneck (Schulz et al. 2005) formations. The Kiscell Clay

Formation, conformably overlying these sediments, contains

a decapod association dominated by Ctenocheles rupeliensis

(Beurlen, 1939) (Beurlen 1939; Hyžný & Dulai 2014;

Fig. 8A, C). From the menilite-type strata of the Carpathian

Foreland Basin near-complete decapod exoskeletons were

reported by Paucă (1929), Jaroš (1939), Jerzmańska (1967)

and Bieńkowska-Wasiluk (2010).

The Transylvanian Basin was a part of the Apuseni shelf

and it was probably connected with the Fore-Rhodopian

Basin of the Tethyan Realm (Popov et al. 2004). In the shal-

low part of the Transylvanian Basin, sandy-calcareous sedi-

mentation took place typified with the Hoia and Mera beds

(Popov et al. 2004). From the Mera Beds numerous ghost

shrimps remains were reported by Bittner (1893).

During the Middle Kiscellian-Solenovian (Late Rupelian),

the entire Paratethys was characterized by brackish salinities

and endemic biota (e.g., Popov et al. 2004; Melinte-

Dobrinescu & Brustur 2008). From this time interval, no

Paratethyan decapod crustaceans are known. In the Late

Oligocene the marine regime was re-established. Marine

connections existed towards the North Sea Basin and the

North Italy Basin via the Slovenian Corridor (Báldi 1986).

Calliax miche lottii is documented in the Oligocene of the

Fig. 8. Selected decapod crustaceans from the Oligocene of the Central Paratethys. A — Ctenocheles rupeliensis (Beurlen, 1939), lectotype

HNHM M.59.4696a. B — Calappilia tridentata (Beurlen, 1939), syntype HNHM M.59.4679. C — Ctenocheles rupeliensis,

HNHM.M.59.4696. D — Loerenthopluma lata Beschin, Busulini, De Angeli & Tessier, 1996, HNHM.M.2010.1.1. Specimens in A– C are

from Óbuda in Budapest (Hungary), specimen in D comes from the borehole Má 115 at Mány (Hungary). Scale bar equals 5 mm. All speci-

mens except B were covered with ammonium chloride prior to photography.

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Northern Lower Miocene of Austria and Slovenia and the

Middle Miocene of Hungary, Slovakia and Slovenia (Hyžný

& Gašparič 2014). This species probably used both seaways

for its spreading. In general, only several decapod species are

known from the Oligocene of the Paratethys (Fig. 3), owing

to the limited exposure of the Oligocene strata and relatively

few published studies on the Oligocene decapods from the

studied area.

Early Miocene: Most of the Early Miocene deep-water

environments were characterized by clayey sedimentation

under dysoxic to anoxic conditions. In the Central Paratethys,

the Eggenburgian is characterized by a transgression with

fully marine conditions as a consequence of an open connec-

tion to the Mediterranean via the Pre-Alpine passage (Sztano

1994). During the Ottnangian, the Alpine Trough was open

(Grunert et al. 2012) and a shallow connection existed

between the Western Paratethys and the North Sea (Martini

1990) (Fig. 1B). In the Central Paratethys, strong Atlantic

influences are observed (Rögl 1999). From the Early Ottnan-

gian of the North Alpine Foreland Basin decapod associations

dominated by Calliax michelottii, Jaxea kuemeli (Fig. 9A)

and Liocarcinus ottnangensis (Bachmayer, 1953a) have been

reported (Hyžný et al. 2015b). By the end of the Ottnangian

a strong regression occurred in the Alpine Fore deep and this

Ottnangian crisis is reflected in brackish water settings with

the endemic Rzehakia fauna (Steininger 1973).

From the Ottnangian (= Helvetian), 10 species were

recorded (Studer 1892, 1898; Bachmayer & Rutsch 1962;

Fraaije et al. 2010 a; Hyžný et al. 2015 b). This relatively low

number probably represents an underestimation influenced

mainly by collection bias. Fine, siliciclastic sedimentation

typical for this stage promotes preservation of near-complete

decapod bodies as exemplified by a mud shrimp J. kuemeli

(Fig. 9A) and a ghost shrimp C. michelottii (Hyžný et al.

2015 b: fig. 5).

In the Karpatian, the marine realm was restricted to the

north-western Pannonian Basin System and the Carpathian

Foreland Basin (Rögl 1999; Mandic et al. 2012). Typical

components of the deep-water habitats (below 200 m) with

muddy bottom are ghost shrimp Callianopsis de Saint Laurent,

1973 (Fig. 9B), squat lobster Munidopsis Whiteaves, 1874

(Hyžný et al. 2014b; Fig. 9C) and brachyuran Styrioplax

Glaessner, 1969. Styrioplax exiguus (Fig. 9E) is known from

the Karpatian of the Slovak part of the Vienna Basin (Hyžný

& Schlögl 2011) and Austrian and Slovenian parts of the

Styrian Basin (Glaessner 1928; Gašparič & Hyžný 2014;

Gašparič & Halásová 2015).

In contrast to undersampled Ottnangian stage, 21 decapod

species were recorded from the Karpatian alone (Fig. 3). This

comparatively higher species richness is a result of recent

efforts to document decapod associations from the Vienna

and Styrian basins (Hyžný & Schlögl 2011; Gašparič &

Hyžný 2015). As taken together, the Ottnangian and

Fig. 9. Selected decapod crustaceans from the Early Miocene of the Western and Central Paratethys. A — Jaxea kuemeli Bachmayer, 1954,

NHMW 2009/0150/0001. B — Callianopsis marianae Hyžný & Schlögl, 2011, KGP-MH CL-011. C — Munidopsis lieskovensis Hyžný &

Schlögl, 2011, KGP-MH CE-005. D — Palaeomyra strouhali (Bachmayer, 1953), holotype NHMW 1953/0051/0001. E — Styrioplax

exiguus (Glaessner, 1928), holotype UMJGP-5453. Specimens in A and D come from Ottnang (Austria), B and C are from Cerová-Lieskové

(Slovakia), and E comes from St. Egidi (Slovenia). Scale bar equals 5 mm. All specimens except C were covered with ammonium chloride

prior to photography.

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Karpatian decapod associations are close to each other not

only taxonomically (Figs. 5, 7) but also in the mode of pre-

servation which is a direct consequence of the prevalent

siliciclastic sedimentation during both stages.

Middle Miocene: In the Central Paratethys, sand-free

calcareous clays (“Tegel facies”) were the most abundant

facies in the Early and Middle Badenian. For this facies, the

most typical decapod species is Tasadia carniolica (Bittner,

1884) (Fig. 10I) which occurs throughout the entire Central

Paratethys. The Early Badenian is characterized by an excep-

tionally rich warm water fauna (Harzhauser & Piller 2007),

including foraminifers, red algae, ostracods, sea urchins,

molluscs, corals, and decapods. Müller (1984a, 1996) and

Collins (2014) reported occurrences of reef-associated deca-

pod faunas from the Early Badenian of Hungary, Poland and

Austria, respectively; typical representatives are Petrolisthes

haydni Müller, 1984a; Kromtitis koberi (Bachmayer &

Toll mann, 1953); Chlorodiella mediterranea (Lőrenthey

in Lőrenthey & Beurlen, 1929) (Fig. 10K); and Daira

speciosa (Reuss, 1871) (Fig. 10J). Nearly a half (47.9 %) of

all Early Badenian decapod species are strictly reef- asso-

ciated (Fig. 3). An even higher proportion (60.9 %) of the

reefal taxa were recorded from the Middle Badenian. One of

the best examples of the Middle Badenian coral patch-reefs

inhabited by decapods is the Fenk quarry in the south-eastern

Vienna Basin (Bachmayer & Tollmann 1953; Riegl & Piller

2000; Müller, 1984a; Hyžný et al. 2014c). Virtually all Middle

Badenian reefal decapod taxa were recorded from this single

locality.

During the Late Badenian, a new transgression began. The

East Slovakian Basin was connected with the Transylvanian

Basin via the Transcarpathian depression and all these basins

were connected with the central Pannonian area (Rögl 1998,

1999; Popov et al. 2004; Fig. 1C). Thriving reef-associated

decapod assemblages are known from the coral settings of

the Great Hungarian Basin (Müller 1984a) and algal- vermetid

reefs of the Carpathian Foreland Basin (Radwański et al.

2006; Studencka & Jasionowski 2011; Górka et al. 2012).

The Late Badenian is still characterized by a warm water

fauna but the overall diversity slightly decreases due to the

onset of the Miocene Climate Transition (Studencka et al.

1998; Kováč et al. 2007; Harzhauser & Piller 2007). The

drop in diversity affects both corals and reef-associated deca-

pod faunas (Müller 1984a: table 2; Fig. 2). In general, how-

ever, Late Badenian is the time of greatest decapod diversity

of the entire Oligo-Miocene of the Central Paratethys

(Fig. 3), partly owing to the collection efforts of Pál Müller

(see above)

The Early Badenian and Late Badenian decapod associa-

tions do not overlap in NMDS ordination (Figs. 5 –7). Taxo-

nomic differences between decapod associations of these two

substages were already demonstrated by Müller (1984a).

Middle Badenian faunas in general are taxonomically much

closer to the Early Badenian ones; this pattern can be

observed in the results of all analyses (Figs. 5 –7).

At the Badenian/Sarmatian boundary, the fossil composi-

tion changed abruptly and stenohaline groups disappeared

(Harzhauser & Piller 2007; Studencka & Jasionowski 2011).

One of the latest marine decapod associations of the Vienna

Basin is documented from the lowermost Sarmatian strata of

the Devínska Kobyla hill (Hyžný 2012; Hyžný & Hudáčková

2012; Hyžný et al. 2012), dominated by ghost shrimps which

are able to tolerate salinity fluctuations.

In the Sarmatian, the open oceanic connections ceased and

the entire Paratethys was inhabited by homogeneous euryha-

line biota, most of which were endemic to the region (Rögl

1998, 1999; Popov et al. 2004). Salinity had regionally

specific composition and ranged from slightly brackish in the

earliest Sarmatian (Studencka & Jasionowski 2011) to hyper-

saline in the Late Sarmatian (Piller & Harzhauser 2005). One

of the Paratethyan endemites was probably also Mioplax

socialis Bittner, 1884, a crab reported from the Badenian

strata of Croatia (Bittner 1884) and the Lower Sarmatian

strata of Austria (Glaessner 1928) and Slovakia (Hyžný &

Ledvák 2014; Hyžný et al. 2016). The euryhaline preferences

of this crab have been demonstrated by Hyžný et al. (2016).

This crab is the only decapod species recorded from the

studied localities from the Sarmatian age suggesting it was

one of the latest marine crabs of the Central Paratethys.

Late Miocene: The Upper Miocene deposits with docu-

mented fauna reflect the existence of a large, long-lived

brackish to freshwater Lake Pannon inside the Carpathian

arch (Fig. 1D) which was separated from the Central Para-

tethys at the Sarmatian/Pannonian boundary (Magyar et al.

1999; Harzhauser & Piller 2007). During that time a shallow

brackish-lacustrine deposition was typical for the Vienna

Basin (Kováč et al. 2004, 2005; Magyar et al. 1999;

Harzhauser et al. 2004). From these deposits fossil burrow

systems attributed to callianassid ghost shrimps were repor-

ted (Hyžný et al. 2015d; Fig. 11H). Later, the Vienna and

Danube basins were filled with deltaic deposits prograding

from the northwest and northeast (Magyar et al. 2007). From

isolated marginal lakes in the west remains of freshwater crabs

were reported (Bachmayer & Pretzmann 1971; Fordinál 1994;

Fordinál & Nagy 1997; Klaus & Gross 2010; Fig. 11A–G).

In the Mediterranean reef-associated decapod faunas still

flourished (Müller 1984b; Gatt & De Angeli 2010).

Reefal versus non-reefal taxa

Ottnangian and Karpatian assemblages differ from the

Badenian ones (Fig. 5). The main contributing factor is the

type of sedimentation and the availability and extent of reef

habitats. Whereas Ottnangian and Karpatian decapod assem-

blages are preserved virtually exclusively in the “Schlier”-

type facies, carbonate sedimentation during the Badenian

and the presence of reefal environments at that time in the

Central Paratethys (Pisera 1996) promoted thriving of more

diverse reef-associated assemblages. As a consequence, the

assemblages from individual basins group in the NMDS

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analysis not according to their spatial proximity but accor-

ding to age. Interestingly, not only Ottnangian-Karpatian

assemblages were distinct from Badenian ones, but Badenian

associations were further subdivided into three distinct

groupings, each for the substages Early, Middle and Late

Badenian (best observable in reefal taxa as shown in Fig. 6).

A great abundance of reefal decapod taxa in the Early and

Middle Badenian is connected to the widespread reefal

Fig. 10. Selected decapod crustaceans from the Middle Miocene of the Central Paratethys. A — Glypturus munieri (Brocchi, 1883), MFGI

M.2355; Rákos, Hungary. B — Eucalliax pseudorakosensis (Lőrenthey in Lőrenthey & Beurlen, 1929), syntype MFGI M.20; Felménes,

Romania. C — Calliaxina chalmasii (Brocchi, 1883), minor chela, MFGI M.21 (holotype of Callianassa rakosensis Lőrenthey, 1897);

Rákos, Hungary. D — Calliaxina chalmasii, major chela, HNHM PAL 2011.33; Rákos, Hungary. E — Pylopagurus leganyi Müller, 1984a,

holotype HNHM 62 3253; Sámsonháza, Hungary. F — Galathea weinfurteri Bachmayer, 1850, NHMW 2013/0580/0014; Deutsch-Alten-

burg, Austria. G — Calappa heberti Brocchi, 1883, MFGI M.2379; Rákos, Hungary. H — Rakosia carupoides Müller, 1984a, NHMW

2015/0052/0007; Gross-Höflein, Austria. I — Tasadia carniolica (Bittner, 1884), MFGI M.35 (holotype of Cancer szontaghii Lőrenthey,

1897); Tasádfő, Romania. J — Daira speciosa (von Reuss, 1871), NHMW 1896/93; Gamlitz, Austria. K — Chlorodiella mediterranea

(Lőrenthey in Lőrenthey & Beurlen, 1929), NHMW 2015/0056/0294; Gross-Höflein, Austria. L — Pilodius vulgaris (Glaessner, 1928),

holotype NHMW 1927/0001/0002; Rauchstallbrunngraben, Austria. M — Xantho moldavicus (Janakevich, 1977), Shepteban, Moldova;

digitalized copy of Janakevich (1969: pl., fig. 6). Scale bar equals 5 mm. All specimens except E, J and M were covered with ammonium

chloride prior to photography.

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settings at that time (Pisera 1996). These settings included

coral patch reefs and coral carpets (Riegl & Piller 2000; Reuter

& Piller 2011), and algal-vermetid reefs (Radwański et al.

2006; Górka et al. 2012). Diverse coral-associated decapod

associations are known from the Lower Badenian strata of

the Styrian, Great Hungarian and Carpathian Foreland basins

(B1-StB, B1-GHB, B1-CFB), the Middle Badenian strata of

the Vienna Basin (B2-VB) and the Upper Badenian strata of

the Great Hungarian Basin (B3-GHB). Decapods associated

with algal-vermetid reefs were reported from the Upper

Bade nian strata of the Carpathian Foreland Basin (B3-CFB).

Palaeobiogeographical implications

Biogeographical homogeneity of Paratethys: The plank-

totrophic larval mode in most anomurans and brachyurans

plays a major role in their dispersal pattern (Gurney 1942;

Moore & McCormick 1969; Harvey et al. 2002, 2014; Martin

2014b). They passively drift in ocean currents promoting

high dispersal potential with larval development having

a great impact on their latitudinal diversity gradients (Fernández

et al. 2009). This is in striking contrast with peracaridans

(e.g., isopods and amphipods) exhibiting direct development

(Harvey et al. 2002; Boyko & Wolff 2014; Wolff 2014) and

consequently restricted dispersal, lower connectivity and

higher potential for isolation. In addition, some decapods are

active swimmers, so the dispersal and migration of decapods

can be very quick (Por 1986). Decapods as palaeobiogeo-

graphical indicators were successfully used in the Northern

Pacific Ocean (Schweitzer 2001) and in the Southern Hemi-

sphere (Feldmann & Schweitzer 2006). Migration pathways

reconstructed in these studies match those observed for mol-

luscs with planktotrophic larval mode earlier. In this study,

the spatial patterns in decapod composition rather demon strate

biogeographical homogeneity of the Paratethys, with sedi-

mentological and palaeoenvironmental conditions being the

major factors influencing distribution of specific taxa.

Already Müller (1984a: 102) recognized that Early, Middle

and Late Badenian reef-associated decapod faunas differ

from each other and proposed several “decapod zones” based

on them. The present results (Fig. 6) corroborate his findings.

The conclusion about the biogeographical homogeneity of the

Paratethyan decapod associations has already been demon-

strated for the Miocene molluscs (Studencka et al. 1998;

Mandic & Steininger 2003) and echinoderms (Kroh 2007).

Rapid dispersal of decapod crustaceans as discussed above

promoted their homogeneous distribution in the Paratethys.

Interactions between Mediterranean and Paratethys:

Previous studies indicate that an alternating anti-estuarine

and estuarine circulation regulated the Middle Miocene faunal

exchange between these two provinces (Báldi 2006: fora-

minifers; Moissette et al. 2006: bryozoans; Harzhauser et al.

2003: molluscs; Kroh & Harzhauser 1999: echinoderms).

For the Early and Middle Badenian, an anti-estuarine

(lagoonal) circulation is assumed, permitting an easier incur-

sion of Mediterranean species into the Paratethys, but hinde-

ring the Paratethyan endemics from entering the Mediterra-

nean. In the Late Badenian, the circulation reversed to estua-

rine type, which is connected with the cooler period (Báldi

2006). As demonstrated by Karami et al. (2011), the Para-

tethys was more responsive to climate change after closure of

the marine seaways than the Mediterranean. Moreover, the

palaeobiogeographic domain formed by the Mediterranean

and the Paratethys was relatively homogeneous during the

Middle Miocene and the similarities with the Atlantic were

smaller (Kroh & Harzhauser 1999; Harzhauser et al. 2003;

Moissette et al. 2006). If this scenario applies to decapod

Fig. 11. Selected decapod crustaceans from the Late Miocene of the Central Paratethys. A — Potamon proavitum Glaessner, 1928, holotype

UMJGP 5828 from Andritz, Austria. B – G — Potamon hegauense Klaus & Gross, 2010, isolated fingers (KGP-MH uncatalogued) from

borehole PiD-1, Slovakia (Danube Basin). H — Fragment of larger complex burrow system Egbellichnus jordidegiberti Hyžný, Šimo &

Starek, 2015 (paratype SNM-Z 37738) attributed to callianassid ghost shrimp (Hyžný et al. 2015d). Scale bar equals 10 mm.

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crustaceans, it will imply that the Middle Miocene Mediter-

ranean and Paratethys decapod assemblages as taken together

were relatively homogeneous.

Gatt & De Angeli (2010: text-fig. 11) compared coral-

associated decapod species from the Middle Miocene (Bade-

nian) of the Paratethys with the association from the Late

Miocene (Messinian) of Malta (Mediterranean): seven species

out of 19 taxa recorded from Malta are shared with the Para-

tethys, whereas similarities are even greater on the genus

level (Müller 1993; Gatt & De Angeli 2010). Based on simi-

larities in decapod faunas and comparison with migration

patterns of other groups, Gatt & De Angeli (2010) suggested

the migration of taxa between the Mediterranean and the

Paratethys during the Langhian. Assuming that the Early and

Middle Badenian exchange flow was controlled by anti-

estuarine circulation (Báldi 2006; Moissette et al. 2006), the

reefal decapod taxa from the Lower and Middle Badenian

strata of the Paratethys are migrants from the Mediterranean.

However, after the reversal in circulation to estuarine type,

the newly evolved Paratethyan species could migrate back to

the Mediterranean. Thus, the Late Miocene reefal decapod

associations could represent (at least partly) descendants of

the Paratethyan migrants.

Paratethys vs. North Sea: Miocene decapod faunas of the

North Sea share numerous genera (and even species) with the

Central Paratethys assemblages. For instance Janssen &

Müller (1984) and Polkowsky (2014) reported a crab Tasadia

carniolica from the Miocene of Belgium and northern Ger-

many, respectively; the species is common in the Middle

Miocene sediments of Austria, Slovakia, Slovenia and even

Romania (Müller 1984a; Hyžný 2011c). Similarly, Fraaije et

al. (2010b) reported another crab, Dromia neogenica Müller,

1978, from the Miocene and Pliocene of the southern North

Sea Basin documenting a link between that basin and the

Central Paratethys and coeval levels in North Africa (Alge-

ria); all these areas share some decapod species. Hyžný &

Gašparič (2014) reviewed all occurrences of a ghost shrimp

Calliax michelottii including those from the Oligocene of

northern Germany and the Miocene of Austria, Hungary, Slo-

vakia and Slovenia representing another example of connec-

tion between the North Sea Basin and the Paratethys. Clearly,

these faunal similarities between the discussed areas are not

an artefact of poor systematics. It is thus reasonable to

assume the migration of decapods sometimes during the

Early Miocene via the Rhine Graben (Martini 1990; Berger

et al. 2005) because no seaway connection existed between

the Central Paratethys and the North Sea during the Middle

Miocene (e.g. Rögl 1998; Harzhauser & Piller 2007). Based

on the proposed estuarine circulation pattern during the Oli-

gocene and Early Miocene (Popov et al. 2004), it is likely

that decapods originated in the Paratethys and migrated into

the Atlantic (North Sea).

Decapods exterminated by the Messinian salinity crisis?

At the end of the Miocene, the Messinian salinity crisis took

place in the Mediterranean basins, thereby also affecting the

Paratethys. This event left marine faunas severely impove-

rished (Hsü et al. 1978; Harzhauser et al. 2002; Krijgsman et

al. 2010; Roveri et al. 2014). Decapod taxa probably

responded differently to the Messinian salinity crisis. For

instance, callianassid genera such as Glypturus and Neocalli

chirus Sakai, 1988, which today are bound to the subtropics

and tropics (e.g. Dworschak & Ott 1993; Felder 2001;

Dworschak 2004), did not survive in the modern Mediterra-

nean. On the other hand, the callianassid genera Calliaxina

Ngoc-Ho, 2003 and Calliax de Saint Laurent, 1973 are still

present in the Mediterranean today (Ngoc-Ho 2003), which

apparently is a consequence of different ecological require-

ments of various ghost shrimp taxa.

A Tethyan origin of Indo-West Pacific faunas? The

simi larities between decapod faunas of the Badenian of the

Central Paratethys and today’s Indo-West Pacific led Müller

(1979a, 1984a) to suggest that Tethyan faunas have their ori-

gin in the Indo-West Pacific. However, he noted that reverse

may also be true (Müller 1979a: 868). The present state of

knowledge of the fossil record suggests migration of Tethyan

marine faunas in an easterly direction during the Oligocene

and/or Miocene. This migration led to a major shift towards

the West Pacific as a centre of diversity, as documented for

molluscs and ophiuroids (Harzhauser et al. 2007, 2008; see

also Renema et al. 2008). Apparently, decapod crustaceans of

the Tethyan stock took the same migratory routes as demon-

strated by Schweitzer (2001), Hyžný (2011a) and Hyžný &

Müller (2012). The fossil record of a mud shrimp Jaxea

Nardo, 1847 suggests its origin in the Tethys before the Early

Miocene. Its descendant lineage is represented by the extant

Jaxea novaezealandiae Wear & Yaldwyn, 1966 (Hyžný

2011a) living in the Western Pacific. The same scenario can

be postulated for a ghost shrimp Glypturus Stimpson, 1866

with its fossil record extending into the Middle Eocene of

Spain and Italy (Hyžný & Müller 2012; Beschin et al. 2012)

and subsequent migration both westwards into the Western

Atlantic and eastward into the Indo-West Pacific (Hyžný &

Müller 2012; Hyžný et al. 2013).

Conclusions

Only a limited number of decapod species are known from

the Oligocene of the Paratethys, owing to the limited expo-

sure of the Oligocene strata and relatively few published

studies on the Oligocene decapods from the studied area.

Whereas Ottnangian and Karpatian decapod assemblages

are preserved virtually exclusively in the “Schlier”-type

facies, carbonate sedimentation during the Badenian and the

presence of reefal environments at that time in the Central

Paratethys promoted thriving of more diverse reef-associated

assemblages.

In all analyses, Early Badenian decapod associations were re-

solved as taxonomically distinct from the Late Badenian ones.

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The Late Badenian was recognized as the time of greatest

decapod diversity of the entire Oligo-Miocene of the Cen-

tral Paratethys, partly owing to collection efforts of Pál

Müller.

Decapod crustaceans of the Paratethys exhibited relatively

homogeneous distribution. The time factor, including specia-

tion and extinction, can explain differences among assem-

blages from the same environment (reefs) but different times.

Based on the distribution of the decapod taxa in the Para-

tethys and adjacent areas, several working hypotheses on

palaeobiogeographical patterns are presented for further

testing: a) the Miocene Paratethyan decapod assemblages

have their origin in the Mediterranean with increasing rate of

endemites due to anti-estuarine migration pattern; b) in the

Miocene the North Sea shared some species with the Para-

tethys suggesting migration via the Rhine Graben into the

Paratethys; c) during the Oligocene and/or Miocene deca-

pods migrated in an easterly direction, thus, contributing to

the modern diversity in the Indo-West Pacific Region.

Acknowledgements: Access to the collections was provided

by Emese Bodor and Klára Palotás (MFGI), Alfréd Dulai

(HNHM), Rok Gašparič (RGA/SMNH), Martin Gross

(UMJGP), Oleg Mandic (NHMW), Barbara Zahradníková

(SNM-Z), and Irene Zorn (GBA). Klement Fordinál (Geo-

logical Institure of Dionýz Štúr, Bratislava) provided mate-

rial from the borehole PiD-1. Mathias Harzhauser, Andreas

Kroh and Oleg Mandic (all NHMW) are thanked for con-

structive comments on the earlier draft of the manuscript.

Oleg Mandic helped with palaeogeographical maps. Thomas

Neubauer (NHMW) and Natália Hudáčková (KGP) are

thanked for help with the NMDS analysis. The manuscript

benefited from the constructive criticism of an associate edi-

tor Adam Tomašových and reviewers Barbara Studencka and

Rafał Nawrot. The research has been supported by the Aus-

trian Science Fund (FWF; Lise Meitner Program M 1544-

B25), VEGA 02/0136/15 and the Slovak Research and

Development Agency under contracts no. APVV-0099-11

and APVV-0436-12.

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