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The events related to the Messinian salinity crisis have been extensively debated since the early 1970s. The spectacular scenario of a completely desiccated Mediterranean subsequently partially occupied by freshwater and brackish endorheic basins triggered a considerable amount of multidisciplinary research for almost five decades. Although the Italian geological record played a crucial role in the origin and complex development of the salinity crisis model, due to the hypothesised palaeobiotic apocalypse, the exploration of the fossil record has been limited or, in certain cases, nearly absent. In this paper, a cursory overview of the Italian fossil record of the Messinian salinity crisis is provided. The integrative analysis of the (primarily) Italian record of microbes, calcareous nannoplankton, dinoflagellates, diatoms, foraminiferans, ostracods, molluscs (and other invertebrates), and fishes reveals the persistence of marine organisms throughout the three stages of the MSC. Moreover, it clearly indicates that a more detailed exploration of the palaeobiological record at a Mediterranean scale is necessary to properly interpret the structure and composition of the biotic communities that inhabited the Mediterranean during the MSC. RIASSUNTO-[La vita in un Mediterraneo completamente disseccato: Uno sguardo al registro paleontologico italiano della crisi di salinità messiniana]-Gli eventi connessi alla crisi di salinità messiniana sono stati profondamente dibattuti sin dall'inizio degli anni 70. Lo spettacolare scenario di un Mediterraneo completamente disseccato e successivamente occupato da bacini endoreici dulcicoli e/o salmastri ha innescato per circa cinque decenni numerose ricerche multidisplinari volte a comprendere le caratteristiche di questo straordinario evento paleoceanografico. Nonostante il registro geologico italiano abbia giocato un ruolo fondamentale nell'origine e nello sviluppo del modello della crisi di salinità, principalmente a causa dell'evocata catastrofe paleobiotica, l'esplorazione del registro paleontologico è stata limitata o, in alcuni casi, del tutto nulla. In questa sede viene fornita una rapida panoramica del registro paleontologico italiano della crisi di salinità messiniana. L'analisi integrata delle evidenze (principalmente) italiane relative a tracce di vita microbica, nannoplancton calcareo, dinoflagellati, diatomee, foraminiferi, ostracodi, molluschi (e altri invertebrati) e pesci suggerisce la persistenza di organismi marini nei tre intervalli della crisi. Inoltre, l'analisi del registro paleontologico indica chiaramente che uno studio maggiormente dettagliato realizzato a scala mediterranea sarebbe necessario al fine di interpretare in maniera più approfondita la struttura e la composizione delle comunità biotiche che caratterizzarono il Mediterraneo durante la crisi di salinità. The great tragedy of Science-the slaying of a beautiful hypothesis by an ugly fact.
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Bollettino della Società Paleontologica Italiana, 58 (1), 2019, 109-140. Modena
ISSN 0375-7633 doi:10.4435/BSPI.2019.04
Living in a deep desiccated Mediterranean Sea:
An overview of the Italian fossil record of the Messinian salinity crisis
Giorgio CA R N E VA L E, Rocco GEN NARI, Francesca LOZAR, Marcello NATALICCHIO, Luca PELLEGRINO &
Francesco DELA PIERRE
G. Carnevale, Dipartimento di Scienze della Terra, Università degli Studi di Torino, Via Valperga Caluso 35, I-10125 Torino, Italy; giorgio.carnevale@unito.it
R. Gennari, Dipartimento di Scienze della Terra, Università degli Studi di Torino, Via Valperga Caluso 35, I-10125 Torino, Italy; rocco.gennari@unito.it
F. Lozar, Dipartimento di Scienze della Terra, Università degli Studi di Torino, Via Valperga Caluso 35, I-10125 Torino, Italy; francesca.lozar@unito.it
M. Natalicchio, Dipartimento di Scienze della Terra, Università degli Studi di Torino, Via Valperga Caluso 35, I-10125 Torino, Italy; marcello.natalicchio@unito.it
L. Pellegrino, Dipartimento di Scienze della Terra, Università degli Studi di Torino, Via Valperga Caluso 35, I-10125 Torino, Italy; lu.pellegrino@unito.it
F. Dela Pierre, Dipartimento di Scienze della Terra, Università degli Studi di Torino, Via Valperga Caluso 35, I-10125 Torino, Italy; francesco.delapierre@unito.it
KEY WORDS - Microbial life, calcareous nannoplankton, dinoagellates, diatoms, foraminiferans, ostracods, molluscs, shes.
ABSTRACT - The events related to the Messinian salinity crisis have been extensively debated since the early 1970s. The spectacular
scenario of a completely desiccated Mediterranean subsequently partially occupied by freshwater and brackish endorheic basins triggered
a considerable amount of multidisciplinary research for almost ve decades. Although the Italian geological record played a crucial role in
the origin and complex development of the salinity crisis model, due to the hypothesised palaeobiotic apocalypse, the exploration of the fossil
record has been limited or, in certain cases, nearly absent. In this paper, a cursory overview of the Italian fossil record of the Messinian salinity
crisis is provided. The integrative analysis of the (primarily) Italian record of microbes, calcareous nannoplankton, dinoagellates, diatoms,
foraminiferans, ostracods, molluscs (and other invertebrates), and shes reveals the persistence of marine organisms throughout the three
stages of the MSC. Moreover, it clearly indicates that a more detailed exploration of the palaeobiological record at a Mediterranean scale
is necessary to properly interpret the structure and composition of the biotic communities that inhabited the Mediterranean during the MSC.
RIASSUNTO - [La vita in un Mediterraneo completamente disseccato: Uno sguardo al registro paleontologico italiano della crisi di
salinità messiniana] - Gli eventi connessi alla crisi di salinità messiniana sono stati profondamente dibattuti sin dall’inizio degli anni 70. Lo
spettacolare scenario di un Mediterraneo completamente disseccato e successivamente occupato da bacini endoreici dulcicoli e/o salmastri
ha innescato per circa cinque decenni numerose ricerche multidisplinari volte a comprendere le caratteristiche di questo straordinario
evento paleoceanograco. Nonostante il registro geologico italiano abbia giocato un ruolo fondamentale nell’origine e nello sviluppo del
modello della crisi di salinità, principalmente a causa dell’evocata catastrofe paleobiotica, l’esplorazione del registro paleontologico è stata
limitata o, in alcuni casi, del tutto nulla. In questa sede viene fornita una rapida panoramica del registro paleontologico italiano della crisi di
salinità messiniana. L’analisi integrata delle evidenze (principalmente) italiane relative a tracce di vita microbica, nannoplancton calcareo,
dinoagellati, diatomee, foraminiferi, ostracodi, molluschi (e altri invertebrati) e pesci suggerisce la persistenza di organismi marini nei tre
intervalli della crisi. Inoltre, l’analisi del registro paleontologico indica chiaramente che uno studio maggiormente dettagliato realizzato
a scala mediterranea sarebbe necessario al ne di interpretare in maniera più approfondita la struttura e la composizione delle comunità
biotiche che caratterizzarono il Mediterraneo durante la crisi di salinità.
The great tragedy of Science - the slaying of a beautiful hypothesis by an ugly fact.
T.H. Huxley
Remarkable hypotheses require extraordinary proof.
R.D. Dietz & M. Woodhouse
FOREWORD: THE DEEP (ITALIAN) ROOTS
OF THE CONCEPT
Almost 50 year after the publication of the famous
paper “Late Miocene desiccation of the Mediterranean” by
Kenneth J. Hsü, William B. F. Ryan and Maria Bianca Cita
(Hsü et al., 1973a) in which the Messinian salinity crisis

scenario that characterised the Mediterranean during
the development of this fascinating event continues to
stimulate considerable interest and cogent debates in the

Deep Sea Drilling Project (DSDP) Leg 13 were crucial to

al., 1973a, b), the concept of a profound environmental
crisis that occurred at a Mediterranean scale at the end of
the Miocene emerged in the 1950s (Denizot, 1952; Selli,
1954; Ogniben, 1957) and was subsequently developed
in the 1960s especially by Raimondo Selli (Selli, 1960)
and Giuliano Ruggieri (e.g., 1962, 1967) based on
onshore studies carried out throughout Italy and Sicily.

considered this Stage as the interval placed between the
Tortonian and the Pliocene (Zanclean), characterised
throughout the Mediterranean by a “crisis of salinity” and
in Italy primarily by evaporitic sediments originated in
hypersaline environments (Fig. 1). Leo Ogniben (1957)
hypothesised that the Messinian evaporitic deposits were
the direct product of the isolation of the Mediterranean
from the Atlantic during the late Miocene. At the same
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Bollettino della Società Paleontologica Italiana, 58 (1), 2019
110

revealed the existence of vast evaporitic deposits beneath

  
et al., 1971; Ryan et al., 1971), recognizing diapiric
structures similar to salt domes that are rooted in a thick
   
Biscaye et al., 1972). The peculiar fossiliferous deposits

investigated by Giuliano Ruggieri who introduced the term
“Lago-mare” co-opting the term “Lac-mer” of Gignoux
     
that characterised the hypothesised endorheic basins
    
waters according to Ruggieri). These basins developed

Miocene, immediately preceding the Pliocene marine

1965).
Since the publication of the results of the DSDP Leg
   
hypothesis of the latest Miocene environmental and
physiographic evolution of the Mediterranean, many
hundreds of papers dealing with the MSC have been
published, exploring all the aspects and consequences
of this event. Due to its spectacular scenario, with the
Mediterranean that was at times completely desiccated,
this event has received considerable attention and publicity
through books, television programs and popular articles
(e.g., Hsü, 1972a, 1984, 2001) and certainly contributed
to the modern establishment of neocatastrophism.
Overall, the role of the Italian sedimentary record (Fig.
1) has been crucial for the origin of the concept and the
development of the knowledge concerning the complex

the Mediterranean during the MSC. The palaeontological
record, however, has been only partially explored and
investigated or, in some cases, totally neglected. The goal
of this paper, therefore, is to provide a cursory overview of
the Italian fossil record of the MSC, restricted to aquatic

understanding of this spectacular event.
OF SALINE GIANTS AND BRACKISH LAKES:
A QUICK LOOK AT THE EVOLVING SCENARIO
OF THE MESSINIAN SALINITY CRISIS
The analysis and interpretation of the data collected
during the Deep Sea Drilling Project (DSDP) Leg 13
led to the birth of the “deep desiccated basin model”
(Hsü et al., 1973a, b), which postulated a Mediterranean
base level laying a few kilometres below the global sea
level and the subsequent development of a giant salt
desert (the “saline giants” of Hsü, 1972b). A primary
consequence of this remarkable catastrophic event was
the annihilation of the resident biota due to the complete
collapse of the Mediterranean marine ecosystem and the
deposition of thick and extensive evaporite successions
in the entire basin, which removed more than 5% of the
dissolved oceanic salts (e.g., Hsü et al., 1977; Rouchy,
    
and oceanographic patterns (e.g., Thunnell et al., 1987).
As a matter of fact the “deep desiccated basin model”
implies that during the MSC, the Mediterranean was
affected by dramatic changes, with the water body
becoming hypersaline, completely desiccated and, then,
hyposaline. The evaporite accumulation was followed by
the deposition of fresh- to brackish water sediments during
the “Lago-mare” event after which the Mediterranean was

Messinian Flooding) with the consequent complete biotic
and environmental recovery of the basin. The progressive
closure of the Atlantic-Mediterranean gateways through

along the African and Iberian continental margins has been
considered as the principal cause that led to the progressive
isolation of the Mediterranean from the Atlantic and the
consequent desiccation of the basin (e.g., Duggen et al.,
2003; Govers et al., 2009).
The adoption of the astronomical cyclostratigraphic
approach and of modern physical stratigraphic concepts
has improved our interpretation of the progression of the
MSC with an accurate and detailed precessional tuning
of the main palaeoenvironmental events. The calibration
of biostratigraphic and geomagnetic Neogene time scale
     
stage at 7.25 Ma (Hilgen et al., 2000) and its end at 5.33
Ma (Van Couvering et al., 2000). Krijgsman et al. (1999)

Fig. 1 - Schematic map of Italy showing the distribution of the

PB: Piedmont Basin; VdG: Vena del Gesso; Tu: Tuscany; Ma:
Marche; LB: Laga Basin; RB: Rossano Basin; BB: Belice Basin;
Cm: Ciminna; CB: Caltanissetta Basin.
111
G. Carnevale et alii - Fossil record of the Messinian salinity crisis
of the MSC in which the onset and development of the
evaporitic phase was constrained between 5.96 and 5.59
Ma and the post-evaporitic phase, characterised by the
deposition of the so-called Upper Evaporites and the
“Lago-mare” sediments, was bracketed between 5.50 and
5.33 Ma. In this context, the cyclostratigraphic calibration
of evaporitic and post-evaporitic sequences seems to result

between 5.59 and 5.50, corresponding to an interval of

of the Mediterranean from the ocean network (Krijgsman
et al., 1999). The desiccation of the Mediterranean during

the MSC characterised by the deposition of non-marine
sediments of the “Lago-mare” event. The remarkable
change of the peri-Mediterranean drainage pattern
produced by the desiccation eventually resulted in the
capture of brackish or fresh waters of Paratethyan origin

lakes and lagoons (e.g., Cita et al., 1978a; McCulloch &
De Deckker, 1989).
Subsequent comprehensive examination of the onshore
sedimentary record throughout the Mediterranean resulted
   
for the MSC (e.g., Clauzon et al., 1996; Roveri et al.,
2001; Rouchy & Caruso, 2006; Manzi et al., 2007),
which culminated with the publication of a consensus
model (CIESM, 2008) based on the strong integration
of bio-, cyclo- and magnetostratigraphic data with
physical stratigraphy and facies analysis. This consensus
stratigraphic model actually provides a new scenario for
the MSC and consists of three main evolutionary stages,
respectively, 1, 2 and 3 (e.g., Manzi et al., 2013; Roveri
et al., 2014a), with stage 3 subdivided into two substages

et al., 2013), evaporites precipitated only in shallow-water
marginal basin where they are represented by a rhythmic
alternation of up to 16 beds of massive selenite (Primary
Lower Gypsum, PLG; e.g., Lugli et al., 2010) and more
or less laminated shales. The formation of gypsum during
this stage was apparently limited to depths shallower than
200 meters, and its deep-water counterparts are dolostones
or, more commonly, organic-rich shales (Manzi et al.,
2007; Lugli et al., 2010; Dela Pierre et al., 2011, 2012;
Natalicchio et al., 2019). The top of the evaporitic deposits

an erosional surface (commonly known as “Messinian
erosional surface”).
The deposits of the second stage (5.6-5.55 Ma) are
grouped into a heterogeneous unit called Resedimented
Lower Gypsum (RLG; Roveri et al., 2008) and are
represented by halite in Sicily, Calabria and Tuscany and,
more commonly, by clastic gypsum deposits (Fig. 1). The
deposits of this stage document the acme of the MSC
with widespread subaerial exposure and erosion possibly
related to a remarkable sea-level drop associated to two
successive glacial stages (TG 14 and TG 12). This interval
of the MSC was likely characterised by a Mediterranean-
scale tectonic activity related to a reorganization of the
Africa-Eurasia plate boundary zone (see Meulenkamp
& Sissingh, 2003). During this stage the exposed PLG
deposits were uplifted, deformed and strongly eroded
and resedimented in deep-water settings producing
clastic gypsum deposits that in many cases encased the
halite deposits together with the calcareous-dolomitic
microbialitic sediments commonly known as “Calcare di
Base” (see Roveri et al., 2014a).
The third stage of the MSC (5.55-5.33 Ma) was
characterised by the deposition of the Upper Evaporites
and the “Lago-mare” event, evidencing an overall

the previous stage. This stage is documented by alternated
evaporites and clastic sediments containing predominantly
  
appear to be widespread in the Mediterranean basin
(see Orszag-Sperber, 2006). The sedimentary sequence
characteristic of this stage of the MSC exhibits a recurrent
vertical organization that allows to separate this stage
     
al., 2014a), between 5.55 and 5.42 Ma, is characterised
by gypsum alternated with shale beds or by shallow- to
deep-water clastic deposits. The fossiliferous content of
the deposits of this substage is generally scarce and the
low values of the 87Sr/86Sr isotope ratio seem to indicate a
substantial freshwater input throughout the Mediterranean.
The second substage (3.2; Roveri et al., 2014a)
started at 5.42 Ma and roughly corresponds with the
“Lago-mare” event. Both the heterogeneous nature
of the sedimentary record and the impossibility of an

the palaeogeographic and palaeoenvironmental context
of the Mediterranean during the third stage of the MSC
(see, e.g., Roveri & Manzi, 2006). For this reason, a
comprehensive interpretation of the “Lago-mare” event
is extremely problematic and the main physiographic
and palaeoecological features at the Mediterranean scale
have been hypothesised based on the analysis of part
of the available fossil record, represented by peculiar
assemblages of brackish or freshwater molluscs and
ostracods traditionally considered of Paratethyan origin
(e.g., Orszag-Sperber, 2006). The palaeoenvironmental
   
indicate that during the third stage of the MSC non-marine
sedimentation took place in the Mediterranean in a series


the basin. The broad distribution of “Lago-mare” deposits
at a Mediterranean scale is interpreted as the product of
the capture of the Paratethyan brackish waters through a
gateway located somewhere in the Aegean (McCulloch
& De Deckker, 1989; Orszag-Sperber et al., 2000) or the
Balkan (e.g., Suc et al., 2015) region, in a general context
    
2002; Cosentino et al., 2005).
       
instantaneous return to fully marine conditions due
     
abrupt collapse of the Gibraltar sill and the origin of
an enormous rapid or gigantic waterfall (McKenzie,
1999; Garcia-Castellanos et al., 2009). This spectacular
event is often recorded by an organic-rich layer of
  
et al., 2008). The magnitude of the sea-level rise that
apparently characterised the Mio-Pliocene transition in
the Mediterranean is not known because of the actual
absence of reliable palaeobathymetric proxies. However,
Bollettino della Società Paleontologica Italiana, 58 (1), 2019
112

with a very limited amplitude of the sea level rise and
a consequent non-catastrophic Messinian-Zanclean
    
Stoica et al., 2016; Krijgsman et al., 2018; Roveri et al.,
2018).
AN OVERVIEW OF THE ITALIAN
PALAEOBIODIVERSITY INVENTORY OF THE
MESSINIAN SALINITY CRISIS
Microbial life
Because the environmental conditions that developed
during the MSC were apparently problematic for a number
of eukaryote groups (e.g., Cita et al., 1978a; Bellanca et al.,
2001; Blanc Valleron et al., 2002), the macro- (and micro-)
fossil content of the Messinian evaporitic units appears to
be scarce and mostly represented by few “extremophilic”

of these group of prokaryotes is therefore crucial to
decipher the chemical and physical conditions in the
water column and in the sedimentary environments during
the course of this dramatic palaeoceanographic event.
In Messinian deposits the evidence of microbial life is
represented by both body and molecular fossils (Vai &
Ricci Lucchi, 1977; Decima et al., 1988; Kenig et al.,
1995; Sinninghe Damsté et al., 1995; Rouchy & Monty,
2000; Guido et al., 2007; Panieri et al., 2010; Turich &
Freeman, 2011; Dela Pierre et al., 2012, 2015; Schopf
et al., 2012; Allwood et al., 2013; Birgel et al., 2014;
Christeleit et al., 2015; Natalicchio et al., 2017; Perri et al.,
2017), although body fossils are especially concentrated

 - Body fossils of prokaryotes are mostly

“Calcare di Base” in Sicily and Calabria (Decima et al.,
1988; Rouchy & Caruso, 2006; Oliveri et al., 2010; Caruso
et al., 2015; Perri et al., 2017) and in the PLG unit in
Northern Apennines and Piedmont. In the PLG unit they
are known from both the bottom grown selenite crystals
(Vai & Ricci Lucchi, 1977; Panieri et al., 2008, 2010;
Schopf et al., 2012; Dela Pierre et al., 2015) and the shales
interbedded to gypsum (Dela Pierre et al., 2014) or, in
certain cases, in the shales representing the deeper water
counterparts of the evaporites (Dela Pierre et al., 2012).

that they considered of algal origin from the carbonatic
layers of the Colombacci Formation, pertaining to the
upper part of the third stage of the MSC, corresponding to

are observed along the vertical growth band of the crystals
(Dela Pierre et al., 2015) and in the re-entrant angle of
the twins (Panieri et al., 2008, 2010; Fig. 2a-b). Such
distribution indicates that the precursor microorganism
lived adhering to crystal faces, thereby suggesting a
benthic lifestyle. In the shales interbedded to gypsum
and in those representing the deep-water equivalents of
 
form dm- thick laminated microbialitic layers (Dela Pierre

formerly described as “spaghetti-like” structures (Vai
& Ricci Lucchi, 1977), consist of curved and straight

rather uniform diameter throughout their lengths (Schopf et
al., 2012; Dela Pierre et al., 2015; Fig. 2b). Well-preserved

by a sequence of rounded cellular compartments of
uniform shape and size. When exposed to UV light the

content (Fig. 2d). Those preserved in gypsum from the
Piedmont basin contain tiny opaque grains within their

and XRD analyses (Dela Pierre et al., 2015). Interestingly,
micro-Raman analyses revealed that these opaque grains
correspond to aggregates of microcrystalline pyrite and,
in rare cases, of polysulphide (Dela Pierre et al., 2015).
     
    
of benthic algae (Vai & Ricci Lucchi, 1977) or of
cyanobacteria (Rouchy & Monty, 2000; Panieri et
al., 2010), suggesting a shallow water depositional
environment located within the photic zone for the
evaporites and associated sediments. This assignment
     
   
bearing gypsum samples from the Vena del Gesso Basin
(Panieri et al., 2010), which is however a controversial
issue (Schopf et al., 2012; Dela Pierre et al., 2015). Other

in the “Calcare di Base”, interpreted these features
either as faecal pellets of brine shrimps (Schreiber,
1978; Natalicchio et al., 2013), indicating shallow and
hypersaline depositional conditions, or of copepods
(Guido et al., 2007), which point to a relatively deep basin


the “Calcare di Base” and the PLG unit have been referred
to as fossils of giant colorless sulphide-oxidizing bacteria
like Beggiatoa and Thioploca (Oliveri et al., 2010; Dela
Pierre et al., 2012, 2014, 2015; Schopf et al., 2012; Perri
et al., 2017). Such an attribution is based on the following
features: 1) absence of terrigenous grains and/or coccoliths
  

that of living giant colorless sulphide-oxidizing bacteria
(e.g., Gallardo, 1977; Fossing et al., 1995; Mussman et
   
80 m in diameter (Schultz & Jørgensen, 2001); 3)
presence of small aggregates of pyrite and associated
polysulphide, which are considered to result from early
diagenetic transformation of original sulphur globules
stored by the sulphur bacteria within their cells (Bailey
et al., 2009; Dela Pierre et al., 2015). These globules
constitute a relevant diagnostic feature for this group of
prokaryotes, representing an intermediate product of the
oxidation of sulphide to sulphate (e.g., Teske & Nelson,
2006). In particular, in modern representatives of the genus
Beggiatoa, polysulphide derives from the rapid (few days)
transformation of cyclooctosulphur (Berg et al., 2014).
 - Despite the study of the
microbial molecular fossils inventory across the entire

revealed the presence of distinct groups of archaea and
113
G. Carnevale et alii - Fossil record of the Messinian salinity crisis
bacteria that thrived during the MSC (Kenig et al., 1995;
Sinninghe Damsté et al., 1995; Turich & Freeman, 2011;
Birgel et al., 2014; Christeleit et al., 2015; Natalicchio
et al., 2017).
Archaeal molecular fossils are among the predominant
lipids found in the pre-MSC and in the MSC deposits
(Birgel et al., 2014; Natalicchio et al., 2017) primarily
consisting in two groups of compounds, glycerol
dibiphytanyl glycerol tetraethers (GDGTs) and diphytanyl
glycerol diethers (DGDs; Fig. 3). GDGTs are particularly
abundant in the pre-MSC deposits as well as in the deposit

show a pattern, dominated by GDGT-0 and crearchaeol
(Fig. 3) similar to that found in modern seawater as well
as in Cenozoic marine sediments (Schouten et al., 2013)
and resembling the GDGT distributions produced by
mesophilic, planktic Thaumarchaeota; these organisms
represent up to 20% of the marine picoplankton in modern
sea water (Wuchter et al., 2006; Schouten et al., 2013),
preferably living in the deeper, meso- to bathypelagic
zones (Karner et al., 2001). The occurrence of marine
planktic archaea suggests the persistence of normal marine
   




Pollenzo section, Piedmont Basin. (a) and (b) are plane-polarised light photomicrographs; (d) is ultraviolet-light photomicrograph.
Bollettino della Società Paleontologica Italiana, 58 (1), 2019
114
conditions in the upper water column at the beginning
of the MSC (Natalicchio et al., 2017; Fig. 3). On the
other side, the DGDs (archaeol and extended archaeol)
provide information about the environmental conditions
at the bottom; these molecular fossils are commonly
produced by extremophilic Archaea (e.g., hypersaline,
methane-rich; Schouten et al., 2013; Birgel et al., 2014)
and extended archaeol, in particular, is only known to be
produced by halophilic Archaea (Dawson et al., 2012).
In the marginal areas of the Piedmont Basin as well as
in the Caltanissetta Basin, these compounds are found to
be especially abundant suggesting the establishment of a
more complex archaeal community at the beginning of the

al., 2014; Natalicchio et al., 2017). Since the conditions
in the upper water column were still favorable for some
algae (see below) and marine archaeal picoplankton (e.g.,
Thaumarchaeota), halophilic Archaea probably inhabited
the water column at, or below, a chemocline as observed
    
et al., 2016).
      
agreement with the occurrence of others peculiar molecular
fossils, such as tetrahymanol (Fig. 3) and his degradation
product gammacerane, in the deposits formed during

et al., 2017) and the Vena del Gesso basins (Kenig et
al., 1995; Sinninghe Damsté et al., 1995; Manzi et al.,
2007). These compounds are common tracers of water
 
Sinninghe Damsté et al., 1995), considered to be sourced
by ciliates (e.g., Harvey & McManus, 1991), anoxygenic

methanotrophs (Banta et al., 2015), which are organisms
commonly living at the chemocline (interface between
oxic and anoxic waters; Wakeham et al., 2007, 2012).
     
Sinninghe Damstè et al. (1995) found the compound

del Gesso Basin; isorenieretane is a lipid synthetised by
anaerobic phototrophic bacteria (Chlorobiaceae), whose
    
within the photic zone.
Finally, bacterial-derived molecular fossils, including
short chain fatty acids (n-C16, n-C18 FA and iso and anteiso-
C15-FA) and hopanoids (bacteriohopanepolyols), are also
important constituent of the “Calcare di Base” in Sicily
and Calabria (Guido et al., 2007; Birgel et al., 2014), as
well as in the deeper counterpart of the PLG units in the
Piedmont Basin (Natalicchio et al., 2017; Fig. 3). These
    
including anaerobic sulfate and iron reducers (e.g., Birgel
et al., 2014; Blumenberg et al., 2015) and aerobic bacteria
(Talbot & Farrimond, 2007).
Calcareous nannoplankton
Calcareous nannofossils are relatively uncommon in
sediments accumulated during the MSC. Their remains
have been reported in Italian onshore sections mainly in

Fig. 3 - Microbial life: molecular fossils. Sketch showing the abundance and distribution of molecular fossils derived from archaea, bacteria


indicated. GDGTs: glycerol dialkyl glycerol tetraethers; DGDs: diphytanyl glycerol diethers; br-GDGTs: branched GDGTs, mostly sourced
by bacteria (Schouten et al., 2013); FA: fatty acids; PLG: Primary Lower Gypsum unit; sterols*: some sterols are possibly sourced also by
terrestrial plants (see Natalicchio et al., 2017 for additional information).
115
G. Carnevale et alii - Fossil record of the Messinian salinity crisis
et al. 2007; Lozar et al., 2010, 2018; Violanti et al., 2013),
although they also occur in those belonging to the second
and third stages of the MSC, namely the laminated gypsum
(Rouchy, 1976; ascribed to the RLG unit by Manzi et al.,
2016) and salt member (Bertini et al., 1998) of Sicily,
as well as in “Lago-mare” deposits (Castradori, 1998;
Cosentino et al., 2012). They have also been recorded in
the deep-sea sediments collected during the DSDP-ODP
cruises (e.g., Sites 132, 134, 653, 654; Roveri et al.,
2014a), but the correlation of these occurrences to one of
the MSC stages is very problematic, due to the complete
absence of reliable biostratigraphic markers. Generally,
the nature of the record of calcareous nannofossils in
the three stages of the MSC is highly debated, since
the salinity crisis paradigm involves a sharp increase in

which would result in the development of a basin not
exploitable by the marine biota. Moreover, during the
third stage of the MSC, they occasionally co-occur with
brackish-water ostracods (see, e.g., Cosentino et al., 2012).
In the sediments preceding the MSC onset the structure
     
restriction of the Mediterranean circulation patterns,
and is dominated by small taxa belonging to the family
Noelaerhabdaceae (Reticulofenestra minuta Roth, 1970
and R. haqi Backman, 1978; Negri & Vigliotti, 1995;
Sprovieri et al., 1996; Negri et al., 1999; Kouwenhoven et
al., 2006; Manzi et al., 2007; Morigi et al., 2007; Iaccarino
et al., 2008; Lozar et al., 2010, 2018; Gennari et al., 2013,
2018a; Violanti et al., 2013) and, to a minor extent, to
the Calcidiscaceae (Calcidiscus spp., Umbilicosphaera
rotula [Kamptner, 1956] and U. jafari Muller, 1974); the
assemblage also includes Helicosphaera carteri (Wallich,
1877), Sphenolithus abies
1954, S. moriformis (Brönnimann & Stradner, 1960), and
Syracosphaera spp. Fluctuations of calcareous nannofossil
relative abundances seem to be correlated to lithological
cyclicity, since during the pre-MSC, the nannofossil

Violanti et al., 2013), with small Reticulofenestra spp.
being very abundant in the low insolation (Maximum
Precession) and Discoaster spp. occurring only in the
maximum insolation (Minimum Precession) parts of
the lithologic cycle (Flores et al., 2005; Violanti et al.,
2013). The Noelaerhabdaceae inhabit the upper portion
of the water column and prefer mesotrophic to eutrophic
conditions, suggesting a well-mixed water column and
high nutrient availability at the surface; they are also
regarded as opportunistic taxa, capable of surviving in
waters with anomalous salinity (Wade & Bown, 2006).
In the euxinic shales deposited in intermediate depths
     

among samples that are otherwise dominated by small
Reticulofenestra (< 3 µm). These consist of S. abies in
the Northern Apennines (Fanantello core; up to 60%
of the total assemblage; Manzi et al., 2007) and of S.
abies, H. carteri, U. rotula and U. jafari in the Piedmont
Basin (Pollenzo, Moncalvo and Banengo sections;
Lozar et al., 2010, 2018; Violanti et al., 2013; Fig. 4a-c).
Peaks of S. abies have been recorded in the sediments
just below the MSC onset in Sicily at Falconara (from
6.45 Ma; Sprovieri et al., 1996; Blanc-Valleron et al.,
     
precessional cycles before the onset of the MSC; Manzi
et al., 2007). These peaks are particularly striking in the
northern Mediterranean records (at Fanantello and in the
Piedmont basin) where this taxon, which never exceeds
12% of the total assemblage in the pre-MSC record,
reaches up to 60% of the total assemblage at the MSC
    
Reticulofenestra (Lozar et al., 2018) in the shale, or by
Umbilicosphaera spp. (Lozar et al., 2018) in the indurated
carbonate beds, are recorded in the Piedmont basin in the
interval interpreted as lateral equivalent of the PLG beds
(Dela Pierre et al., 2011; Fig. 4c-d). In the Fanantello core
S. abies peaks have been recorded in
the sediments equivalent to the lower PLG cycles (Manzi
et al., 2007).
Peaks of S. abies have been also observed in the
laminated gypsum of two Sicilian sections, Montedoro and
Siculiana, probably belonging to the second stage of the
MSC (Rouchy, 1982; Rouchy & Caruso, 2006), together
  U. jafari, U.rotula,
Cd. cfr. leptoporus, R. minuta. Additional calcareous
nannofossil assemblages are recorded in the clayey
layers in the Racalmuto salt mine (Bertini et al., 1998),
with small Noelaerhabdaceae and Coccolithus pelagicus
(Wallich, 1877), Sphenolithus spp., U. rotula, Calcidiscus
spp., Pontosphaera spp., Rhabdosphaera spp., H. carteri,
Discoaster quinqueramus Gartner, 1969, Amaurolithus
primus (Bukry & Percival, 1971), A. delicatus Gartner &
Bukry, 1975, Syracosphaera spp., as well as in the shales
intercalated with the gypsum in the Crostolo River section
in the Northern Apennines (Barbieri & Rio, 1974).
In the marls intercalated with the laminated gypsum
in Sicily (balatino-like gypsum, belonging to the second
stage of the MSC), Braarudosphaera spp. have also been
found (Schreiber, 1974), suggesting brackish conditions
in the upper water column as also reported by stable
isotope analyses (Pierre, 1974). In the Caltanissetta Basin

assemblages have been described in the laminated
gypsum, both in the pelitic laminae (Cd. leptoporus
[Murray & Blackman, 1898], S. abies, Scyphosphaera
apsteinii Lohmann, 1902, R. pseudoumbilicus [Gartner,
1967]) and the calcareous laminae (dominated by small
Reticulofenestra spp.; Rouchy, 1982).
In the onshore record of the “Lago-mare” event
(Cosentino et al., 2006, Mondragone well; Cosentino
et al., 2012, Fonte dei Pulcini), coccoliths have been
interpreted as reworked, due to the high abundance of
reworked Cretaceous species, together with Palaeogene
and long ranging taxa (Reticulofenestra spp., Sphenolithus
spp., Coccolithus spp.). This interpretation has been
reconsidered by Pellen et al. (2017), who reported the
occurrence of Ceratolithus acutus Gartner & Bukry, 1974
in the same Fonte dei Pulcini section, thereby suggesting

in uppermost Messinian sediments; the co-occurrence
     
    
during the third stage of the MSC has been hypothesised
by Crescenti et al. (2002) for the S. Nicolao section.
     
Tyrrhenian sites 132, 653B and 654A calcareous
Bollettino della Società Paleontologica Italiana, 58 (1), 2019
116
nannofossils have been reported from the evaporitic unit
(Site 132, cores 21-27; Site 654A, cores 28-35; Müller,
1990) or just above it (Site 653B, core 25; Pierre &
Rouchy, 1990). In the sediments overlying the evaporitic

the presence of dwarfed foraminiferans has also been
reported. The ooze interbedded in the halite in Site 134
(cores 9 and 10) contains an oligotypic assemblage with
D.
challengeri Bramlette & Riedel, 1954 and D. variabilis
Martini & Bramlette, 1963. In Site 654A (cores 26 to
36) Müller (1990) recorded calcareous nannofossils
(of smaller size than normal) co-occuring with gypsum
and dolomite. The assemblages consist mainly of C.
pelagicus, R. pseudoumbilicus, H. carteri, Cd. leptoporus,
S. abies, U. rotula, Cd. macintyrei (Bukry & Bramlette,
1969), and Pontosphaera multipora (Kamptner, 1948 ex
 A. delicatus
and discoasterids are also present.
Dinoagellates
Studies on late Miocene dinoflagellate cysts are
primarily focused on the assemblages recorded in MSC-
related deposits. The late Tortonian and early Messinian
    
conditions with abundant outer platform and oceanic
taxa (e.g., Santarelli et al., 1998; Londeix et al., 2007). At
least in the Caltanissetta Basin, the sediments pertaining

diversity of dinocyst species that provides evidence of
considerable hydrological mixing in the basin suggesting
the coexistence of taxa typical of confined neritic
environments (Homotryblium spp. and Lingulodinium
machaerophorum
indicative of oceanic marine biotopes with normal salinity
and normal to sub-normal hydrology (Impaginidium spp.,
Nematosphaeropsis labyrinthus [Ostenfeld, 1903] and
Spiniferites mirabilis [Rossignol, 1964]). According to
Londeix et al. (2007), these data suggest a Mediterranean
Fig. 4 - Calcareous nannoplankton. Light microscope images from the PLG unit, Pollenzo section, Piedmont Basin. a) Abundance peak of
Sphenolitus abies (thin arrow), together with Umbilicosphaera rotula (thick arrow) and Helicosphaera carteri (empty arrow); scale bar: 10
Sphenolitus abies
Umbilicosphaera rotula (possible coccosphere). d) Tiny Umbilicosphaera spp. and Syracosphaera
117
G. Carnevale et alii - Fossil record of the Messinian salinity crisis
   
stage of the MSC.
Diverse assemblages of dinoflagellate cysts have
also been reported from the clayey layers within the
halite sequences of the Realmonte and, especially, the
Racalmuto salt mines, thereby providing information
about the diversity and abundance of these planktonic
algae during the second stage of the MSC (Bertini et
al., 1998). The taxa recognised in these assemblages
are typical of warm to warm-temperate surface waters.
Overall, these assemblages mostly consist of neritic
(Lingulodinium machaerophorum, Operculodinium
israelianum [Rossignol, 1962], Polysphaeridium zoharyi
[Rossignol, 1962]) and oceanic (Impaginidium sp.,
Nematosphaeropsis labyrinthus, N. lemniscata [Bujak,
1984], Spiniferites hyperacanthus
1955], S. mirabilis, S. ramosus [Ehrenberg, 1838]) taxa
that indicate the presence of normal marine conditions in
the basin during the deposition of the clayey sediments
(Bertini et al., 1998).

is rather sparse in sediments documenting the substage 3.1
and relatively abundant in those accumulated during the
“Lago-mare” event (substage 3.2). In the upper portion of
the Upper Evaporites of the Caltanissetta Basin, Londeix et

reworked assemblages dominated by taxa indicative of
Homotryblium
plectilum [Drugg & Loeblich, 1967]) associated with less
abundant open marine oceanic taxa (Impaginidium spp.).
On the other side, the dinocyst assemblages of the “Lago-
mare” event have been extensively debated (e.g., Londeix
et al., 2007; Popescu et al., 2007, 2009; Pellen et al., 2017;
Grothe et al., 2018) probably because of their problematic
interpretation. As far as concerns the Apennine record,
based on the palaeoecological (and palaeobiogeographic)
 
al. (2017) evidenced a complex scenario with alternated
marine and brackish episodes, of which the latter
were characterised by a relevant contingent of taxa of
  
Formation in Sicily, Londeix et al. (2007) recognised a
remarkable dominance of euryhaline dinocysts with a
subordinate amount of open marine taxa. In the context

taxon Galeacysta etrusca
(as well as its often associated Pyxidinopsis psilata [Wall
& Dale in Wall et al., 1973] and Spiniferites cruciformis
Wall & Dale in Wall et al., 1973), commonly regarded
as a palaeoenvironmental and palaeobiogeographic
marker for the “Lago-mare” event thought to be a
Paratethyan immigrant. The biogeographic history of
this taxon has been recently summarised by Grothe et al.
(2018). According to these authors (Grothe et al., 2018),
Galeacysta etrusca appeared in the Pannonian basin at
about 8 Ma and subsequently dispersed into the Dacic,
Black Sea and Caspian basins at about 6.1 Ma following
the establishment of physical connections between them.
    
apparently occurred in the Mediterranean during the
  
Bertini, 2006; Londeix et al., 2007; Iaccarino et al., 2008;
Cosentino et al., 2012; Pellen et al., 2017), documenting
the existence of physical and ecological conditions that
allowed the dispersal of this Paratethyan taxon into the
Mediterranean between 5.37 and 5.33 Ma.
Diatoms
The Messinian record of diatoms (Bacillariophyceae)
is mostly restricted to the pre-MSC successions (~7-6
Ma), where diatomaceous sediments occur regularly
alternated to sapropels and marls (Pellegrino et al., 2018).

MSC are scattered and consistently thinner (e.g., Müller
& Schrader, 1989; Fourtanier et al., 1991). The limited
accumulation of diatom-rich sediments during the MSC
has been usually explained with the establishment of
hypersaline settings that precluded the proliferation of
diatoms (e.g., Selli, 1954).
This interpretation discouraged a systematic
investigation of the Messinian evaporitic units aimed to the

this reason, the majority of researches concerning the late
to latest Messinian diatom record of the Mediterranean,
are focused on the scattered, intra-evaporitic diatomaceous
layers and on deep-sea, gypsum-free sediments indicative
of a variety of brackish to fully marine environmental
conditions (e.g., Schrader & Gersonde, 1978; Müller &
Schrader, 1989; Fourtanier et al., 1991).
Recently, however, well-preserved diatoms have been
observed in selenitic gypsum (Dela Pierre et al., 2015;
     
Galeacysta etrusca

Bollettino della Società Paleontologica Italiana, 58 (1), 2019
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in the Piedmont Basin (Banengo section). The diatoms
occur intermingled to brownish-greenish organic remains
   
   
belong to extant families, genera or informal groups
with well-known general ecological habits (Round et al.,
1990). Among them, Chaetoceros sp. vegetative frustules

remains (Fig. 7a-e), together with Biddulphia sp., this
latter often occurring as nicely preserved long chains
(Fig. 6a-e). Naviculoid diatoms have been recorded,
but their small size did not allow a precise generic and

belonging to the genus Grammathophora sp. have been
also observed (Fig. 6g), together with rare specimens of
Stephanodiscaceae, Surirella sp. and Rhizosolenia sp.
(Figs 6i, l and 7f).
The consistently good preservation of the observed
specimens, especially the delicate vegetative frustules
of Chaetoceros sp. with joined setae, allows to rule
out the possibility of reworking, at least for the most
representative component of the assemblage. The epipelic
and epiphytic diatoms were instead reasonably transported

their relatively good preservation, the transport was
probably limited.
It is interesting to note the abundant occurrence of
 Chaetoceros sp.
vegetative frustules in the examined samples. Diatom
blooms are typically followed by biological or physical
aggregation, favoured by the production of sticky
biopolymers (TEP) and by the physical entanglement
of frustules, which promote the rapid sink of diatom
aggregates, preserving them from dissolution and
zooplankton grazing (e.g., Passow et al., 1994; Passow,
2002). However, once reached the sediment-water
interface, the diatom frustules must be rapidly buried in
order to be preserved. Therefore, the fast-growing gypsum
crystals may have favoured the rapid entombment of the
frustules.
These diatom assemblages, characterised by the co-
occurrence of abundant Chaetoceros sp. remains mixed
with planktic, epipelic and epiphytic, marine to freshwater
diatoms, do not support a hypersaline setting linked to a
strongly evaporated water column, but rather a coastal

The overall low amount of diatom specimens in the
Messinian gypsum-bearing sediments can be considered
following two perspectives: it can be a genuine by-product
of limited diatom productivity or, alternatively, the result
of a selective preservation of some specimens. In the

instead of hypersalinity, other triggering factors should
be considered in order to interpret the supposed low rates
of opaline productivity during the late Messinian (e.g.,
reduced bioavailability of limiting factors such as Si,
P, N and Fe). In the second case, conditions favourable
to opal production in the upper water column may
have co-occurred with the establishment of a chemical
environment unfavourable to opal preservation on the
sea-bottom. The possible implications of the second
scenario are intriguing, especially considering the ability
of microbial consortia involved in the sulfur cycle to
modulate the pH of pore waters (e.g., Gallagher et al.,
2012; Pace et al., 2017), a crucial parameter controlling
silica solubility (e.g., Ehrlich et al., 2010). However, only
a wider, more detailed morphological analysis, coupled to
biomarker investigation of gypsum-rich units, may shed
light on this hypothesis.
The deposits of the third stage of the MSC, both the
Upper Evaporites and those recording the “Lago-mare”
event are rarely characterised by the presence of a diatom

(1978) documented poorly diversified assemblages
with shallow water euryhaline taxa, suggesting salinity

Foraminiferans
The foraminiferans record a progressive increase of
stressed conditions in the upper water column and at the


both planktonic and benthic organisms (Blanc-Valleron et
al., 2002; Sierro et al., 2003; Kouwenhoven et al., 2006;
    
as, since the beginning of the Messinian, their absence in

the Falconara section) and the disappearance of oxyphilic
taxa at intermediate depths (Fig. 8; Kouwenhoven et al.,
2003; Iaccarino et al., 2008) seem to be indicative of a
severe oxygen depletion.
A further step at 6.7 Ma marks the increasingly
oligotypic character of the assemblages, which show
a typical precession-driven imprint (e.g., Sierro et
al., 2003). This pattern is given by the dominance of
warm/oligotrophic planktonic and absence of benthic
foraminiferans in the sapropels (tuned to insolation
maxima) and cold/eutrophic planktonic with abundant
benthic in marls or diatomite (insolation minima). From
6.4 Ma, benthic assemblages are dominated by buliminids
and bolivinids (Fig. 8) indicating high organic carbon

salinity and/or low oxygen content (Kouwenhoven et al.,
2003).
As far as concerns the planktonic taxa, the decrease in
diversity is evidenced by the disappearance of the deep
dwelling globorotalids at around 6.7 Ma (Globorotalia
scitula 
and 6.5 Ma (Globorotalia miotumida Jenkins, 1960 and
Globorotalia conomiozea Kennett, 1966, which are
included in the G. miotumida gr.) (Fig. 8). Furthermore,
stressed upper water column conditions are well depicted
by the occurrence of the endemic Mediterranean
Turborotalita multiloba (Romeo, 1965) ranging from
Fig. 6 - Diatoms. The epipelic-epiphytic-planktic marine inshore to freshwater diatom assemblage from the PLG unit, Banengo quarry, Piedmont
Basin. a-e) BiddulphiaBiddulphia sp. and Grammatophora
Grammatophora
Surirella
119
G. Carnevale et alii - Fossil record of the Messinian salinity crisis
Bollettino della Società Paleontologica Italiana, 58 (1), 2019
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6.415 to ca. 6.05 Ma (Fig. 8). This morphospecies
is thought to derive from Turborotalita quinqueloba
(Natland, 1938) and adapted to highly eutrophic (Sierro et
al., 2003) or highly saline surface waters (Blanc-Valleron
et al., 2002).
The progressive stressed conditions culminate at
the onset of the MSC, which is usually marked by the
complete disappearance or, at least, by a strong decrease
of either benthic or planktonic foraminiferans with a size
> 125 µm. In most central and peripheral Mediterranean
basins this event well approximates the MSC onset
(Manzi et al., 2007, 2013, 2018; Gennari et al., 2013,
2018a; Violanti et al., 2013). This fact is reported in the
biostratigraphic zonation of the Mediterranean with the
Non Distinctive Zone, which corresponds to the MSC
time interval (5.97-5.33 Ma; Iaccarino et al., 2007). The
apparent disappearance of the marine microfauna at the
onset of evaporite deposition was one of the evidences
that led to the desiccation scenario (Hsü et al., 1973b).
More recently, the foraminifer characterization of MSC
deposits has been considerably improved. In particular,
it has been noted that smaller foraminiferans (mainly T.
quinqueloba and Globigerina bulloides d’Orbigny, 1826
in the 45-125 µm size fraction; Fig. 8) survived and are

stage of the MSC (e.g., Violanti et al., 2013; Corbì &
Soria, 2016; Manzi et al., 2018).
According to Catalano et al. (2016), a different
scenario appears to be recorded in Sicily where the barren
interval extends back before the MSC onset and also
  
the disappearance of calcareous microfossils seems to be

origin in more marginal setting. The environmental
perturbation causing the disappearance of foraminiferans
seems to be diachronous (Bellanca et al., 2001), primarily
related to the proximity to the coast. A strong reduction
in abundance of foraminiferans also predates the onset of

section, Cyprus Island (Eastern Mediterranean) (Gennari
et al., 2018a). This reduction follows a tectonic pulse
which could had promoted increasing terrigenous input
18O
values and 87Sr/86Sr ratios deviating from oceanic values
suggest that the freshening of the upper water column was
the main trigger of foraminiferan decrease.
Data from the deep settings of the central portion of the

MSC. In the Levantine and Tyrrhenian basins the PLG unit
seems to be devoid of foraminiferans, which apparently
disappeared at the onset of the MSC (Roveri et al., 2014b;
Manzi et al., 2018). Studies addressing the foraminiferan
     
concentrated in the westernmost Mediterranean area.
In the Sorbas and Almeria-Nijar basins several authors
observed that foraminiferans are still present in the
pelitic intercalations between the gypsum layers of the

the occurrence of foraminiferans within the pelitic
intercalation of the Yesares Fm. (PLG unit) in the Nijar

to those of the pre-evaporitic sediments and include both
benthic (Ammonia sp., Elphidium sp., Bolivina plicatella
Cushman, 1930, Bolivina spathulata [Williamson,
1858], Bulimina aculeata d’Orbigny, 1826, Cassidulina
spp., Cibicidoides pseudoungerianus [Cushman, 1922],
Uvigerina sp., small epiphytes, miliolids and “fragile
nodosariids”) and planktonic (G. conomiozea, G. scitula,
Neogloboquadrina acostaensis [Blow, 1959], both
sinistral and dextral, T. quinqueloba and small-sized
globigerinids) taxa. However, the strictly marine taxa are
considered to be reworked and only inner shelf euryhaline
taxa or those thriving in shallow lagoonal environments
are regarded as autochthonous (Fig. 8). The occurrence of
G. conomiozea and of sinistral N. acostaensis is considered
as the main evidence of reworking since N. acostaensis
was prevalently dextral in both the Mediterranean and the
Atlantic from 6.34 Ma whereas G. conomiozea is reported
to be extinct in the Mediterranean before the onset of the
MSC (Fig. 8; Van de Poel, 1992). In the same area, Riding
 
sized, marine assemblage in the upper part of the Yesares
Fm. (PLG unit) and in the overlying Sorbas Member.
These assemblages are considered as in situ; however,

the two formations. At an equivalent stratigraphic position
(top of the PLG unit), Aguirre & Sánchez-Almazo (2004)
documented a marine foraminiferan assemblage, yielding
G. miotumida, which is not regarded to be extinct before
the onset of the MSC by the authors. Goubert et al. (2001)
collected samples in the infra-gypsum intercalations of

of the MSC (PLG unit) in the marginal (western) portion
of the Sorbas Basin. They observed that planktonic
foraminiferans are very rare (only Trilobatus trilobus
[Reuss, 1850] was observed) in the marls between the
third and fourth gypsum layer. Ammonia tepida (Cushman,

and second cycles of the PLG unit; in the upper part of
the section Porosononion granosum (d’Orbigny, 1846)
became dominant, thereby suggesting an upward transition
from an infralittoral to inner circalittoral environment. In
the Bajo-Segura Basin, Corbì & Soria (2016a) reported
dwarf-foraminiferans in the marly intervals intercalated
to the gypsum layers of the PLG unit. These small-sized
assemblages also yield dextral N. acostaensis and G.
miotumida gr. specimens. The small globigerinids are
interpreted as the result of stressed marine condition in
     
Ammonia and Elphidium individuals.
The sedimentary products of the second stage of the
MSC are mainly restricted to the central and deep part of
the Mediterranean where few data are available. The salt
mines of Sicily are among the very few places where clay
intercalation within the halite deposits can be examined.
Bertini et al. (1998) examined nine of these intercalations

foraminifer assemblages in three levels (P/B ratio is 90-
Fig. 7 - Diatoms. The planktic marine diatom assemblage from the PLG unit, Banengo quarry, Piedmont Basin. a-e) Chaetoceros sp., vegetative
frustules. f) Rhizosolenia
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G. Carnevale et alii - Fossil record of the Messinian salinity crisis
95%). Below and above these three levels, the samples
exhibit very low diversity and abundance (scattered T.
quinqueloba and Globigerina sp.) or are completely
barren. The intermediate fossiliferous samples yielded both
long-range taxa (Globigerinoides spp., Globoquadrina
spp. and Orbulina spp.) and a numbers of Late Neogene
globorotalids (both keeled and not-keeled). The latter are
represented by sinistral Globorotalia menardii (Parker,
Jones & Brady, 1865 after d’Orbigny, 1826 nomen
nudum), G. miotumida gr., Globorotalia suterae Catalano
& Sprovieri, 1971, Globorotalia saphoae Bizon & Bizon,
1965 and Globorotalia sphaericomiozea Walters, 1965;
N. acostaensis is also present with predominantly dextral
specimens. Although rather rare, benthic foraminiferans

The third stage of the MSC is the most
controversial in term of foraminifer characterization
and palaeoenvironmental significance. The benthic
euryhaline Ammonia tepida or A. beccarii (Linnaeus,
1758) are generally documented as the most abundant
and widespread taxa. They occur in variable abundance
in successions outcropping in peripheral areas, but also
in DSDP-ODP holes in the more central sectors of the
Mediterranean, where they are usually less common.
Ammonia
intercalated to the selenites of the Upper Evaporites in
Sicily (Bonaduce & Sgarrella, 1999; Grossi et al., 2015)
and in Cyprus (Orszag-Sperber, 2006) or in equivalent
selenite-free deposits underlying the Messinian/Zanclean
boundary (Hole 968A, Blanc-Valleron et al., 1998; sites
375-376, sites 965 to 968, Orszag-Sperber, 2006; Bajo-
Segura Basin, SE Spain, Soria et al., 2008a). Ammonia
often co-occurs with ostracods of the genus Cyprideis; this
biofacies apparently represents a good biostratigraphic
marker of the lower part of the third stage (3.1; Iaccarino
& Bossio, 1999; Grossi & Gennari, 2008). Ammonia can
be associated to other shallow benthic taxa, such as the
euryhaline Elphidium sp. (Site 967, Spezzaferri et al.,

include rare Cribroelphidium sp., Haynesina germanica
(Ehrenberg, 1840), Ammobaculites sp. and Neoconorbina
sp. (e.g., Trave section, Northern Apennines; Iaccarino
et al., 2008). In a few cases, the infralittoral Nonion
boueanum (d’Orbigny, 1846) can co-occur with Elphidium
(Nijar Basin, Bassetti et al., 2006; Montepetra borehole,
Northern Apennines, Grossi & Gennari, 2008). In the
Garruchal Fm. of the Bajo-Segura Basin, Soria et al.
(2008b) also documented a lagoonal palaeoenvironment
with the occurrence of the miliolid Quinqueloculina
laevigata d’Orbigny, 1826 together with Ammonia,
Elphidium spp. and H. germanica.
However, a diversified planktonic and/or benthic

in some cases associated with the shallow benthic taxa
    
deposits were postulated (Cita et al., 1978a, b) during the
examination of the material collected during the DSDP
drillings in the 1970s. In some cases, the planktonic stock
can be considered clearly reworked from older rocks since
it contains a mixture of Cretaceous, Eocene, Oligocene
or early/middle Miocene taxa. Clear examples are those
of the Eraclea Minoa section (Sicily, Italy; Roveri et al.,
2006) and Polemi Basin (Cyprus; Orszag-Sperber et al.,
2006) and Site 967 (Spezzaferri et al., 1998). Instead, the
occurrence of Messinian biostratigraphic markers like
the G. miotumida gr. together with long range Neogene

to Aguirre & Sánchez-Almazo (2004), the deposits of
the third stage of the MSC of the Nijar Basin record
the alternation of marine shelf setting, characterised
by marine pelagic assemblages, and continental/deltaic
setting, characterised by brackish benthic foraminiferans

Bassetti et al. (2006); based on the recognition of G.
menardii gr. 4, other than G. miotumida and the mixing
of shallow and deep-water foraminiferans, the authors
considered the assemblages of the third stage of the Nijar
Basin as entirely reworked. In the Sorbas Basin, the Sorbas
     
stage of the MSC) and record the third stage in a more
marginal setting (Fortuin & Krijgsman, 2003; Roveri et
al., 2019), characterised by the absence of foraminiferans,
except a thin level yielding an almost monogeneric
bolivinid assemblage (Gennari et al., 2018b). According
to Clauzon et al. (2015), the Zorreras Mb. should be
attributed to the Zanclean also based on the occurrence
of Globorotalia margaritae Bolli & Bermudez, 1965 and
Sphaeroidinellopsis specimens in a thin clay layer at the
Sorbas/Zorreras transition. An assemblage composed of
Bolivina cf. paralica Perconig, 1952, Ammonia tepida,
Rosalina sp. and small globigerinids is reported by
Iaccarino & Bossio (1999) at the Messinian/Zanclean
transition of the Balearic Rise (ODP Site 975). The
deposits of the third stage of the MSC are often proposed
to host marine foraminifers heralding the Zanclean full
restoration of marine condition. A transitional fauna
is reported in Hole 969B (Eastern Mediterranean;
Spezzaferri et al., 1998), Montepetra borehole (Northern
Apennine; Grossi & Gennari, 2008), Hole 975 (Balearic
Rise; Iaccarino & Bossio, 1999; Iaccarino et al., 1999),
and Kalamaki (Zakynthos Island, Karakitsios et al., 2017).
These assemblages are all very similar to those found in
Fig. 8 - Foraminiferans. Schematic characterization of the foraminifer assemblages of the MSC in the Mediterranean area. a-j) Planktonic
foraminifera: a) Globorotalia menardii; b) Globorotalia miotumida gr.; c) Globorotalia scitula; d-e) Neogloboquadrina acostaensis sinistral
and dextral coiling, respectively; f) Turborotalita multiloba; g) idealised small globigerinid assemblage; h) ideal basal Zanclean assemblage
composed of the long range taxa Trilobatus trilobus, Globigerina bulloides, Globoturborotalita decoraperta (Takayanagi & Saito, 1962),
Turborotalita quinqueloba, Globigerinita glutinata (Egger, 1893), Neogloboquadrina acostaensis
in the lowermost two precessional cycles, respectively); i) Sphaeroidinellopsis seminulina (Schwager, 1866); j) Globorotalia margaritae.
k-w) Benthic foraminiferans: k) Siphonina reticulataCibicidoides italicus (Di Napoli, 1952); m) Uvigerina cylindrica
gaudryinoides Lipparini, 1932; n) Hanzawaia boueana (d’Orbigny, 1846); o) Bulimina aculeata; p) Bolivina dilatata Reuss, 1850; q) Bolivina
spathulata; r) Ammonia tepida; s) Porosononion granosum; t) Elphidium macellum (Fichtel & Moll, 1798); u) Haynesina germanica; v) Nonion
boueanum; w) A. tepida, Bolivina cf. paralica and Rosalina
(2000), Milker & Schmiedl (2012), Corbì & Soria (2016) and http://www.mikrotax.org/pforams/index.php?dir=pf_cenozoic.
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the basal Zanclean and dominated by planktonic genera,
such as Globigerinoides, Globigerina, Globoturborotalita,
Trilobatus and Globigerinita (Iaccarino et al., 1999);

Almázo, 2004), in these transitional assemblages the
G. miotumida gr. is absent and N. acostaensis shows a
prevalently dextral coiling.
Ostracods
Ostracods are among the most extensively studied fossils

the palaeogeographic and palaeoenvironmental features
of the third stage. Decima (1964), Ruggieri & Greco
      
recognised the common occurrence of brackish ostracod
taxa in the peculiar upper Messinian deposits overlying
the evaporites in Sicily and Northern Apennines. An
extensive exploration of Messinian deposits throughout
the Mediterranean area evidenced a substantial change of
the ostracod assemblages with the apparent disappearance
of the fully marine and diverse pre-MSC taxa (see, e.g.,
Benson, 1976; Sissingh, 1976; Benson et al., 1991). The

rarely provided ostracod remains (see, e.g., Decima,
1964), whereas abundant and sometimes diverse
ostracofaunas are well known from the deposits of the
third stage, especially from those documenting the “Lago-
mare” event (e.g., Gliozzi, 1999; Grossi et al., 2008).
The ostracod assemblages typical of the “Lago-mare”
sediments are characterised by Paratethyan taxa that
dispersed into the Mediterranean as a result of the capture

the MSC (e.g., McCulloch & De Deckker, 1989; Bonaduce
& Sgarrella, 1999; Orszag-Sperber, 2006). Although some
authors considered the typical “Lago-mare” ostracod
taxa as Mediterranean endemics (Bassetti et al., 2003),
      
undeniable (e.g., Gliozzi et al., 2007; Stoica et al., 2016).
Carbonnel (1978) defined the Loxoconcha djafarovi
Zone to emphasise the biostratigraphic relevance of the
Paratethyan immigrants in constraining the “Lago-mare”
event. More recently, the biostratigraphic hypothesis
     
Grossi et al. (2011) who provided a new scheme with
Fig. 9 - Ostracods. a-j) Schematic distribution of selected taxa in the second and third stages of the MSC following the biozones proposed by
Grossi et al. (2011). a) Loxoconcha mülleri (Mehes, 1908); b) Tyrrhenocythere pontica (Livental in Agalarova et al., 1961); c) Loxoconcha
eichwaldi; d) Cyprideis agrigentina Decima, 1964; e) Loxocorniculina djafarovi; f) Euxinocythere praebaquana; g) Amnicythere propinqua;
h) Caspiocypris pontica (Sokac, 1972); i) Tyrrhenocythere ruggierii Devoto in Colacicchi, Devoto & Praturlon, 1967. Scale bar: 0.1 mm.

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G. Carnevale et alii - Fossil record of the Messinian salinity crisis
two biozones, the Loxoconcha mülleri Biozone spanning
from 5.59 to 5.40 Ma and the Loxocorniculina djafarovi
Biozone, which covers the uppermost portion of the third
stage of the MSC from 5.40 to 5.33 Ma (Fig. 9). Both

species. The effectiveness of this ostracod-based
biostratigraphy is rather problematic since the appearance
of the Paratethyan taxa throughout the Mediterranean is
possibly diachronous (see, e.g., Carbonnel, 1980), as also
suggested by the occurrence of typical “Lago-mare” taxa
Amnicythere propinqua [Livental,
1929], Euxinocythere praebaquana [Livental in Agalarova
et al., 1940], Loxocauda limata [Schneider in Agalarova
et al., 1940], Loxoconcha eichwaldi Livental, 1929,
Loxocorniculina djafarovi [Schneider in Suzin, 1956],
Zalanyiella venusta [Zalanyi, 1929]) in deposits clearly

Marmolaio; Caponi, 2008).
The Italian late Miocene fossil record includes
numerous brackish ostracod assemblages, many of which
preceding the “Lago-mare” event (Gliozzi et al., 2005,
2007; Faranda et al., 2007; Ligios et al., 2012; Colombero
et al., 2014). These assemblages are currently well known
and provide a good opportunity to interpret the structure
and composition of the ostracod faunas characteristic
      
MSC. Although the taxonomic composition of the late
Tortonian and early Messinian ostracod assemblages is
    
faunas, several brackish taxa (genera or species) of
clear Paratethyan affinity (Amnicythere, Bakunella,
Camptocypria, Chartocythere, Labiatocandona,
Lineocypris, Loxoconchissa, Mediocytherideis,
Propontoniella) can be documented in Italy at least
since the Tortonian (e.g., Gliozzi et al., 2007; Ligios
      
are also documented in late Serravallian deposits of
the Ebro Basin, Spain (Gliozzi et al., 2005, 2007).
According to Gliozzi et al. (2007), the presence of
    
Tortonian or early Messinian should be related to episodic
passive dispersal events via aquatic birds, whereas the
successive diverse “Lago-mare” contingent represents
the unambiguous evidence of the establishment of direct
geographic connections that allowed an active dispersal
from the Paratethys into the Mediterranean during the
final stage of the MSC. The possibility of episodic
passive dispersal via aquatic birds is evocated due to
the lack of evidence for a direct connection between the
Mediterranean and Paratethys before the “Lago-mare”
event. However, although passive dispersal via aquatic
birds is a well-known dispersal strategy for freshwater
ostracods characterised by parthenogenetic reproduction
or for sexually reproducing species in which females
exhibit internal brood care (e.g., Whatley, 1990, 1992),
the majority of the hemicytherids, leptocytherids and
loxoconchids genera (likely including those reported
in late Tortonian and early Messinian Italian deposits)
reproduces sexually and not retains the fertilised eggs
within the carapace (see, e.g., Boomer et al., 1996),
making them not particularly susceptible to transport
by aquatic birds. Consequently, their arrival in the
Mediterranean necessarily implies the existence of one or
more direct brackish connection(s), at least temporarily
open, that allowed the immigration from the Paratethys.
Such connection(s) possibly promoted the arrival of
the typical “Lago-mare” taxa at least in the complex
palaeogeographic context that occurred in the hinterland
     
stage of the MSC, whereas the humid climatic phase and
widespread development of marginal and satellite basins
throughout the Mediterranean during the “Lago-mare”

remarkable demographic explosion of these Paratethyan
immigrants (see Carnevale et al., 2018).
Molluscs (and other invertebrates)
Together with the ostracods, molluscs are among
the most extensively studied fossils from MSC-related
deposits, particularly those of the third stage documenting
the “Lago-mare” event.
Just before the onset of the MSC a diverse marine
mollusc fauna inhabited the Mediterranean (e.g.,
Compagnoni, 1964; Ruggieri et al., 1969). According to
   

interval just preceding the onset of the MSC exhibited a
Mio-Pliocene Atlantic-Proto-Mediterranean distribution,
whereas the other taxa can be referred to as Miocene
Proto-Mediterranean endemics, some of which became
extinct at the end of the Miocene. Assemblages of fully
marine infralittoral and circalittoral molluscs are known

Italy and Spain (e.g., Bossio et al., 1978; Montenat et al.,
1980; Lacour et al., 2002; Néraudeau et al., 2002), and
provide a further evidence to the persistence of marine
conditions during this part of the crisis. The fossiliferous

the Sorbas Basin and also contain remains of bryozoans,
polychaete tubes, decapod crustaceans and abundant sea
urchins (e.g., Montenat et al., 1980; Néraudeau et al.,
2001; Lacour & Néraudeau, 2002). Bryozoan remains
are also known from the clayey strata intercalated to the
selenitic gypsum layers at Cava Marmolaio in Tuscany
(Caponi, 2008).
Brackish and freshwater molluscs are well-known
from Messinian deposits (e.g., Esu & Girotti, 1989;
Ligios et al., 2012), especially from those documenting
the “Lago-mare” event. The mollusc assemblages
characteristic of the “Lago-mare” event are known from
several localities in Italy (and Sicily), as well in other
Peri-Mediterranean sectors. As far as the Italian localities
are concerned, these peculiar mollusc assemblages have
been reported since the end of the XIX century (e.g.,
Capellini, 1879, 1880; Sacco, 1886). The “Lago-mare”
mollusc assemblages exhibit a peculiar composition with
abundant bivalves of the genus Dreissena and the cardiid
subfamily Lymnocardiinae and a variety of continental
and brackish gastropods (Fig. 10), including the very
common Melanoides, Melanopsis, Saccoia and Theodoxus
(e.g., Esu & Girotti, 1989; Harzhauser et al., 2015). These
assemblages are usually indicative of oligo- and meso-
haline waters and have been traditionally considered as
the product of a massive dispersal from the Paratethyan
basins into the Mediterranean during the latest part of the
MSC (e.g., Orszag-Sperber, 2006; Esu, 2007). However,
Bollettino della Società Paleontologica Italiana, 58 (1), 2019
126
recent studies have revealed that some presumed “Lago-
mare” taxa occurred in the Mediterranean well before the
MSC (e.g., Ligios et al., 2012), and most of them actually
represents Mediterranean endemics (Esu & Popov,
2012; Harzhauser et al., 2015; Colombero et al., 2017).
Moreover, the analysis of the geochemical signature of
these molluscs indicates that the paralic palaeobiotopes

with thalassogenic waters (Grunert et al., 2016).
Fishes and other vertebrates

Sicilian deposits originated during the MSC, representing
a largely unexploited source of palaeoenvironmental
information about the faunal and ecological structure
of the Messinian aquatic palaeobiotopes. As a matter
of fact, the (often neglected) relevance of fishes in
palaeoenvironmental studies lies in their bio-ecological
characteristics; because of their mobility and migratory
behaviour, fishes can provide valuable information
about a vast array of contiguous biotopes, whereas their
trophic level attribution is unambiguously indicative of
the relative size and complexity of the aquatic food web.
These bio-ecological features can be extremely useful to
contribute to the interpretation of the palaeogeographic
and palaeoenvironmental context of the Mediterranean at
least during the intervals of the MSC for which these are

resolution and/or extremely heterogenous nature of the
sedimentary products.
    
both neritic and oceanic, are known from the Messinian
deposits predating the onset of the MSC (e.g., Landini &
Menesini, 1984; Gaudant, 2002; Carnevale, 2003, 2004,
2006; Carnevale & Bannikov, 2006; Carnevale & Pietsch,
2006). The palaeoichthyological record is particularly rich
in Italy (e.g., Sturani & Sampò, 1973; Bradley & Landini,
1984; Bedini et al., 1986) and Sicily (e.g., Arambourg,
1925; Leonardi, 1959; Gaudant et al., 1996), where

sapropels or diatomites, which accumulated (everywhere
in the Mediterranean) during the pre-MSC interval of the
Messinian stage in response to the precessional forcing of


dates back to earliest part of the XIX century when the
Fig. 10 - Molluscs. a-g) Gastropods and bivalves from the “Lago-mare” deposits of Moncucco Torinese, Piedmont Basin: a) Theodoxus
mutinensis (D’Ancona, 1869); b) Melanoides curvicosta (Deshayes, 1835); c) Melanopsis narzolina d’Archiac in Viquesnel, 1846; d) Saccoia
oryza Brusina, 1893; e) Euxinicardium subodessae (Sinzov, 1877); f) Pontalmyra bollenensis (Mayer, 1871); g) Dreissena ex gr. rostriformis

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G. Carnevale et alii - Fossil record of the Messinian salinity crisis
prominent Swiss naturalist Louis Agassiz (1832) described
the cyprinodontid Aphanius (= Lebias) crassicaudus
(Agassiz, 1832) based on material collected from the

outcropping near Senigallia. Since that time a considerable
amount of Messinian palaeoichthyological data has
been accumulated and abundant fossils of Aphanius
crassicaudus have been documented for the three stages
of the MSC in all the sectors of the Mediterranean (Fig.
11). For this reason, this extinct cyprinodontid species is
commonly regarded as an icon of the MSC palaeontology
(e.g., Gaudant, 1979; Sorbini & Tirapelle Rancan, 1979;
Gaudant et al., 1988; Landini & Sorbini, 1989; Carnevale
et al., 2018).
A taxonomic and ecologically heterogenous fish
     
Oreochromis lorenzoi,
Borgo Tossignano (PLG unit), Northern Apennines, scale bar: 10 mm; b) Mugil cf. cephalus Linnaeus, 1758, Cava Serredi (“Lago-mare”
deposits), Tuscany, scale bar: 10 mm; c) Lichia  amia, left cleithrum, Borgo Tossignano (PLG unit), Northern Apennines, scale bar:
50 mm; d, Aphanius crassicaudus, left lateral view, Pollenzo section (RLG unit), Piedmont Basin, scale bar: 10 mm; e) Diaphus befralai
Brzobohaty & Nolf, 2000, Cava Serredi (“Lago-mare” deposits), Tuscany, scale bar: 1 mm; f) Diaphus splendidus (Brauer, 1904), Podere
Torricella (“Lago-mare” deposits), Tuscany, scale bar: 1 mm; g) Myctophum tchi (Schwarzhans, 1979), Podere Torricella (“Lago-mare”
deposits), Tuscany, scale bar: 1 mm; h) Hoplostethus cf. mediterraneus, Moncucco Torinese (“Lago-mare” deposits), Piedmont Basin, scale
bar: 1 mm; i) Diaphusrubus Girone, Nolf & Cavallo, 2010, Moncucco Torinese (“Lago-mare” deposits), Piedmont Basin, scale bar:
1 mm; j) Gadiculus labiatus, Podere Torricella (“Lago-mare” deposits), Tuscany, scale bar: 1 mm; k) Physiculus sp., Moncucco Torinese
(“Lago-mare” deposits), Piedmont Basin, scale bar: 1 mm.
Bollettino della Società Paleontologica Italiana, 58 (1), 2019
128
(Fig. 11), documented by a number of localities in the
Piedmont basin (Castagnito, Cherasco, Costigliole d’Asti,
Guarene d’Alba, Monticello d’Alba, Piobesi d’Alba, Santa
Vittoria d’Alba, Scaparoni), Northern Apennines (Borgo
Tossignano, Brisighella, Monte delle Formiche, Monte
Faeti, Pietralacroce, San Lazzaro di Savena, Senigallia,
Sirolo) and Tuscany (Cava Marmolaio, Cava Migliarino).
From an ecological point of view, this assemblage is
dominated by marine (e.g., Lepidopus spp., Lichia
amia [Linnaeus, 1758], Microchirus abropteryx [Sauvage,
1870], Mugil sp., Sarda sp., Sardina sp., Scorpaena cf.
minima Kramberger, 1882, Spratelloides sp., Trachurus
spp., Thunnini indet.) and estuarine (Aphanius spp.,
Atherina spp., Atherinomorus? etruscus [Gaudant, 1978],
Gobius spp.) taxa, although a few freshwater (Lates
cf. niloticus Linnaeus, 1758, Oreochromis lorenzoi
Carnevale, Sorbini & Landini, 2003, Paleoleuciscus cf.
oeningensis [Agassiz, 1832], Salvelinus oliveroi Gaudant
in Cavallo & Gaudant, 1987) and diadromous (Alosa
crassa Sauvage, 1873, Clupeonella maccagnoi Gaudant
in Cavallo & Gaudant, 1987) species are also present (e.g.,
Sturani, 1973; Gaudant, 1981; Cavallo & Gaudant, 1987;
Landini & Sorbini, 1989; Carnevale et al., 2003, 2008a).
Fish remains are reported from the deposits of
the second stage of the MSC in the Piedmont Basin
(Carbonara Scrivia), Northern Apennines (Cajariccia,
Camignone, Lunano, Monte Sant’Angelo, Monte
Castellaro) and Tuscany (Saline di Volterra), and in
the Caltanissetta Basin in Sicily (Aragona, Canicattì,
Castrogiovanni, Feudo Muscini, Montedoro, Portella di
Pietro, Realmonte, San Cataldo, Solfara Casino, Solfara
di Palagonia) and are usually represented by monotypic
assemblages with abundant remains of the estuarine
cyprinodontid Aphanius crassicaudus (e.g., D’Erasmo,
1928; Sorbini & Tirapelle Rancan, 1979; Gaudant et al.,
1988; Landini & Sorbini, 1989; Fig. 11). The marine
round herring Spratelloides lemonei Arambourg, 1927
is also relatively common in these deposits. The most

is that of Monte Castellaro (Sorbini, 1988; Landini &
Sorbini, 1989) that includes a variety of marine neritic
(Epinephelus sp., Harengula sp., Microchirus abropteryx,
Spratelloides sp., Zeus primaevus Cocchi in Massalongo
& Scarabelli, 1859) and oceanic (Capros arambourgi
Baciu, Bannikov & Santini, 2005, Maurolicus muelleri
[Gmelin, 1789]) taxa together with some rare freshwater
euryhaline (Lates niloticus, Oreochromis lorenzoi) and
estuarine (Aphanius crassicaudus, Atherina boyeri Risso,
1810) species.
Carnevale et al. (2018) recently summarised the
available data about the ichthyofaunal assemblage of the
       
remains are primarily represented by otoliths, which are
known from at least six localities (Cava Serredi, Capanne
di Bronzo, Ciabot Cagna, Podere Torricella, Moncucco
Torinese, Le Vicenne) recording the “Lago-mare” event
(Carnevale et al., 2006a, b, 2008b, 2018; Colombero et
al., 2017); a single locality pertaining to the substage 3.1,

remains (Colombero et al., 2014). About 50 species-level
taxa are known from this stage, among which marine
neritic (e.g., Aphia minuta Risso, 1810, Argyrosomus sp.,
Batrachoididae indet., Blennius sp., Grammonus sp., Liza
sp., Pagellus sp., Spratelloides sp., Sprattus sp., Umbrina
sp.) and oceanic (Benthosema spp., Bolinichthys italicus
[Anfossi & Mosna, 1971], Ceratoscopelus sp., Diaphus
spp., Gadiculus labiatus [Schubert, 1905], Hoplostethus
cf. mediterraneus Cuvier in Cuvier & Valenciennes,
1829, Hygophum spp., Lampadena gracile [Schubert,
1912], Myctophum coppa Girone, Nolf & Cavallo, 2010,
Physiculus

(Leptosciaena caputoi Bannikov, Schwarzhans &
Carnevale, 2018, Trewasciaena kokeni [Schubert, 1902])

with the Paratethyan basins (e.g., Bannikov et al., 2018).
    

mare” event) seems to provide unambiguous evidence
of the presence of normal marine conditions in the
Mediterranean before the Messinian-Zanclean boundary,
demonstrating that a new interpretation of the “Lago-
mare” event is necessary.
Overall, the palaeoichthyological record of the MSC
indicates a remarkable degree of ecological homogeneity
throughout the three stages of the crisis, with a nearly
continuous presence of marine steno- and euryhaline taxa.
Such a faunal continuity seems to be suggested also by
the comparative analysis of the taxonomic composition of
the late Miocene (pre-MSC) and Zanclean ichthyofaunas
(see Carnevale et al., 2018).

of the MSC, cranial remains of a whale have been recently
found in the Terminal Carbonate Complex of the island of
Mallorca (Mas et al., 2018). Despite the age of Terminal
Carbonate Complex is rather controversial (e.g., Roveri et
al., 2009), Mas et al. (2018) suggested a correlation with
the second stage of the MSC.
SYNTHESIS
The picture emerging from the integrative analysis of
the data provided herein reveals that the palaeontological
record of the MSC (Fig. 12) is not fully complete for

the same time, it is certainly not inadequate to contribute
to the interpretation of the patterns of palaeoenvironmental
evolution of the Mediterranean between 5.97 and 5.33
Ma. To date, only a limited role has been attributed to the
fossil record in the characterization of the environmental
scenario of the MSC (e.g., Roveri et al., 2014a). The
evocated incompleteness of the record itself related to the
catastrophic biotic annihilation, and the apparent peculiar
composition of certain fossil assemblages have been
used as key arguments for excluding a large part of the
potentially available palaeontological information from
the discussion about the palaeoenvironmental evolution of
the Mediterranean. The apparent inadequacy of the fossil
documentation is certainly related to the nature of the

the widespread development of “stressed” environmental
(and depositional and taphonomic) conditions at least

development of unfavourable environmental conditions
in the water column began before the onset of the MSC
129
G. Carnevale et alii - Fossil record of the Messinian salinity crisis
and eventually resulted in the apparent extinction of
the euhaline benthic biota as well as of the calcareous
plankton, with the exception of the small-sized foraminifer
assemblages that seems to survive. During the past decades,
the absence of these components of the aquatic ecosystem
was roughly interpreted as the palaeobiological evidence
of the catastrophic hydrological and geomorphological

In concrete terms, because of the catastrophic scenario
evocated for the MSC, the sudden disappearance of
euhaline benthos and calcareous plankton was implicitly
considered as the product of the ecological collapse of
the Mediterranean marine biome and of the complete
annihilation of the aquatic biota. This approach, however,
is based on the assumption that the palaeontological record
is always reasonably complete and that the absence of
record necessarily corresponds to the reliable record of
the (original) absence. Moreover, as mentioned above, the
negative palaeobiological evidence used to support the
catastrophic scenario did not include all the components
of the original biota potentially available in the record and
in part reported herein.
The cursory survey of the Italian fossil record of
      
    
information and, consequently, to remark the necessity
to properly include the fossil record in the (still) cogent
debate about the MSC. The palaeontological data discussed
above suggest an almost continuous presence of aquatic
organisms throughout the MSC (Fig. 12), implying that
complex and heterogenous aquatic biotopes persistently
occurred between 5.97 and 5.33 Ma. This conclusion
is in large part consistent with the modern views about
the palaeoenvironmental evolution during the MSC
(e.g., Roveri et al., 2014a), which are mostly based on
stratigraphic, sedimentological and geochemical data and
suggest that a waterbody was present in the Mediterranean
at least for most of the crisis. The chemical nature and
structure of the waterbody, as well as its cyclic variation

of the waterbody has been recently demonstrated for the

c), the chemical features of the Mediterranean waters for
the third stage are still poorly understood. The benthic
assemblages with ostracods and molluscs have been
used to postulate the “Lago-mare” scenario with the
    
waters of Paratethyan origin (e.g., Cita et al., 1978a), despite

of the groups of fossils discussed in the text in the Italian fossil record. See text for a detailed explanation. Solid line indicates common
occurrence while dashed line indicates episodic or rare occurrence. APTS: astronomical polarity time scale; m.f.: molecular fossils.
Bollettino della Società Paleontologica Italiana, 58 (1), 2019
130
the limited knowledge of the palaeobiological features of
these taxa. The recurrent benthic assemblages characterised
  
“Lago-mare” biofacies actually represent local indicators
of marginal shallow and brackish water conditions. The
typical “Lago-mare” benthic assemblages are generally
found in marginal or satellite basins (e.g., Gliozzi, 1999),
which largely developed during the Messinian, as a result
   
tectonic deformation of the Neogene (Roveri et al., 2001)
     
These assemblages have been occasionally reported also
in a few of the many deep-sea sites drilled, where they are
probably reworked from the marginal basins (Riding et
al., 1998). As mentioned above, because of their mobility

about a vast array of contiguous biotopes, in this case about
those present in the open marine area scarcely documented
in the onshore sedimentary record. Fish remains suggest
that heterogeneous and highly diverse marine biotic
communities were present during the third stage of the
MSC, providing a clear evidence that the Mediterranean
never transformed into a brackish lake system (Carnevale
et al., 2018). The calcareous plankton (nannoplankton
and foraminiferans) that often occurs in the deposits of
the third stage (in the whole Mediterranean), especially
those recording the “Lago-mare” event, is commonly
considered as reworked from older rocks, or, at the very
least, as evidence of short-living marine ingressions (e.g.,
Spezzaferri et al., 1998; Iaccarino & Bossio, 1999; Iaccarino
et al., 1999; Rouchy et al., 2001). In a number of cases,
there is no evidence of reworking, due to the lack of the
typical features such as size sorting or mixing of species of

 
the ecologically diverse ichthyofaunas and their associated
calcareous plankton necessarily imply the re-establishment
of the whole marine biota and their complex ecological
intrarelationships (see Carnevale et al., 2006b).
Summarizing this long discussion, it is reasonable
to conclude that the persistence of marine organisms
throughout the three stages of the MSC is indicative of
the persistence of a marine Mediterranean during this
crucial interval of the Cenozoic history. Therefore, the
   
of the Mediterranean in some ways similar to the so-
called “deep-water deep-basin” hypothesis postulated
by Schmalz (1969, 1991) and corroborated by the ideas
of Debenedetti (1982) and Roveri et al. (2014c). In this
context, the peculiar sedimentary products of the MSC
should be regarded, at least in part, as the result of the


Atlantic gateways, tectonic activity, climate and eustasy
(see Roveri et al., 2014c).
FUTURE DIRECTIONS
The considerable amount of papers dedicated to the
      
of the spectacular scenario evocated for this late
Cenozoic event. Several aspects of this breath-taking
event have been explored in great detail also outside
    

by the MSC model that has represented an apparently
remarkable explanatory mechanism for the present
disjunct geographical distribution of continental peri-
Mediterranean organisms (e.g., Bocquet et al., 1978;
Bernini, 1984; Bianco, 1990; Zardoya & Doadrio, 1999).
Together with the considerable media promotion, the vast
use of the model by biologists contributed to strengthen the
plausibility of this hypothesis as a remarkable geological
discovery that has achieved textbook stature (e.g., Stanley,
1989). Paradoxically, despite a number of stratigraphical,
sedimentological, geodynamical, geochemical and
geophysical (etc.) investigations have been devoted to
characterise the MSC, the contribution of palaeontology
has been comparatively limited and primarily focused at
supporting the catastrophic scenario of an “oceanographic
apocalypse” (e.g., Taviani, 2002) at the end of the
Miocene. The survey of the Italian fossil record presented
herein, although far from being exhaustive, demonstrates
that the palaeontological information potentially available
is abundant and qualitatively adequate to contribute
to the discussion. The analysis of the record suggests
that a relevant environmental perturbation certainly
took place in the Mediterranean between 5.97 and 5.33
Ma but, at the same time, clearly indicates that marine
organisms persisted throughout the three stages of the
MSC. However, a more detailed exploration of the fossil
record of the MSC at Mediterranean scale is necessary
to expand our knowledge about the structure and
composition of the Mediterranean biotic communities.
Moreover, a comprehensive comparative examination of
the pre-MSC and Zanclean Mediterranean and Eastern
Atlantic (Morocco, Portugal, Spain) fossil record would
be crucial to properly interpret the biotic continuity vs
turnover across the MSC (see Neraudeau et al., 2001) and
to evaluate the plausibility of the western Mediterranean
“sanctuaries” or “refugia” that allowed the Pliocene
survival of the Miocene endemics (e.g., Grecchi, 1978;
David & Pouyet, 1984; Moisette & Pouyet, 1987; Di
Geronimo, 1990).
ACKNOWLEDGEMENTS
We are particularly obliged to Richard W. Jordan (Department
of Earth and Environmental Sciences, Yamagata University,
Yamagata) for useful comments and discussion about diatom
  
constructive suggestions for its improvement, we are particularly
grateful to Konstantina Agiadi (Department of Historical Geology
and Palaeontology, National and Kapodistrian University, Athens),
Walter Landini (Dipartimento di Scienze della Terra, Università di

Sciences, Prague), and an anonymous reviewer. The research was
supported by grants (ex-60% 2016-2017 and 2018) to G.C. of the
Università degli Studi di Torino.
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Manuscript received 12 February 2019
Revised manuscript accepted 20 March 2019
Published online 30 April 2019
Guest Editors Massimo Bernardi & Giorgio Carnevale
... The accumulation of huge amounts of evaporitic deposits that occurred during the first phase of the salinity crisis was followed by the capture of Eastern Pararethyan non-marine waters that partially refilled the basin leading to the deposition of the fresh-or brackish-water sediments of the Lago-Mare event (e.g., Cita et al. 1978;McCulloch & De Deckker 1989). The adoption of such a scenario necessarily implies an oceanographic apocalypse in the Mediterranean (see Taviani 2002), with the annihilation of its marine eukaryotic biota and the collapse of the marine ecosystem, followed by the establishment of brackish taxa of Paratethyan origin during the Lago-Mare event and the subsequent complete biotic and environmental recovery after the basal Zanclean re-flooding (see Carnevale et al. 2019). ...
... Although this detailed stratigraphic model clarifies several aspects of the progression of this complex event, a comprehensive definition of the paleoenvironmental scenarios remains largely elusive, as well as the nature and extent of the involved sea level changes. The fossil content of the evaporitic and associated deposits has been discussed by Dela Pierre et al. (2015), Carnevale et al. (2019) and Pellegrino et al. (2021) and was shown to be rich and diverse although this has often been underestimated. Conversely, considerable efforts were devoted to the peculiar latest Messinian ostracod and mollusk assemblages of Paratethyan affinity traditionally used to define the environmental context of the Lago-Mare event. ...
... At Kanthikas, fish remains have been observed in the laminated marls below the fifth gypsum bed (between 5.44 and 5.42 Ma) of the Upper Gypsum unit (Rouchy et al. 2001), while at Lethymbou they have been discovered in the marls underlying the second gypsum bed (between 5.52 and 5.50 Ma) of the Upper Gypsum unit (Manzi et al. 2016) the MarIne paleoIchthyologIcal record of the Msc The skeletal record Articulated skeletal remains of teleost fishes are common fossils in MSC-related deposits, especially in those recording the first two stages. As a matter of fact, one of the paleontological icons of the MSC is a teleost fish, the cyprinodontid Aphanius crassicaudus (Agassiz, 1832), which was widespread in the Mediterranean before and throughout the crisis almost always represented by articulated skeletons, often showing pachyostotic condition (Gaudant 1978(Gaudant , 1979Sorbini & Tirapelle Rancan 1980;Gaudant et al. 1988;Landini & Sorbini 1989;Carnevale et al. 2018Carnevale et al. , 2019. ...
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Almost fifty years after the first definition of the Messinian salinity crisis (MSC), the events that occurred in the Mediterranean during the terminal portion of the Miocene still attract the attention of a large and diverse scientific community. Although fossils are relatively common in the deposits that accumulated during the MSC, their significance for the interpretation of the latest Miocene paleoenvironmental evolution of the Mediterranean has been underevalued. In this paper, we summarize the marine paleoichthyological record of the three stages of the MSC based on both articulated and isolated skeletal remains and otoliths, the latter almost exclusively known from the Lago-Mare phase. We focus on the composition of the marine ichthyofauna of the Mediterranean during the three main stages of the MSC, showing the persistent continuity of marine stenohaline taxa throughout most of the interval between 5.97 and 5.33 Ma. While the record of articulated fish skeletons is unquestionably autochthonous, thereby providing unambiguous evidence of the occurrence of open marine environments in the MSC preceding the Lago-Mare phase, the autochthonous nature of the otolith record has often been questioned. For this reason, the otolith record of marine fishes has been examined in detail from a taxonomic and paleoecological point of view. Three species, Bellottia verecunda n. sp., Benthosema taurinense n. sp., and Bostrychus marsilii n. sp., are described as new and a thorough discussion about the possible origin of the otoliths is provided. Alternative explanations for the occurrence of otoliths of marine fish during the Lago-Mare phase, such as reworking, contamination from overlying Pliocene sediments or import from outside the Mediterranean through aquatic birds are considered unlikely. In our assessment, the occurrence of marine fish otoliths in the Lago-Mare phase can be explained with the presence of normal marine environments in the Mediterranean, at least temporarily. Therefore, we suggest that the paleoichthyological data provided herein should be integrated in the future evolutionary paleoenvironmental reconstructions of the MSC.
... Warren, 2016;Benison, 2019). Microorganisms have been isolated from salt crystals, evaporite deposits and associated fluid inclusions, in the rock record (Stan-Lotter et al., 2002;Vreeland et al., 2007;Schubert et al., 2010;Fendrihan et al., 2012;Carnevale et al., 2019). For example, great palaeobiodiversity is described for the Miocenic Messinian evaporitic deposits, Italy (Carnevale et al., 2019). ...
... Microorganisms have been isolated from salt crystals, evaporite deposits and associated fluid inclusions, in the rock record (Stan-Lotter et al., 2002;Vreeland et al., 2007;Schubert et al., 2010;Fendrihan et al., 2012;Carnevale et al., 2019). For example, great palaeobiodiversity is described for the Miocenic Messinian evaporitic deposits, Italy (Carnevale et al., 2019). Recently, the unique extreme environments at the Dallol geothermal area, Ethiopia Cavalazzi and Filippidou, 2021) have provided the opportunity to study life at its biophysical limits in a hypersaline poly-extreme environment (Belilla et al., 2019;Gómez et al., 2019). ...
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In this work, we address the difficulty of reliably identifying traces of life on Mars. Several independent lines of evidence are required to build a compelling body of proof. In particular, we underline the importance of correctly interpreting the geological and mineralogical context of the sites to be explored for the presence of biosignatures. We use as examples to illustrate this, ALH84001 (where knowledge of the geological context was very limited) and other terrestrial deposits, for which this could be properly established. We also discuss promising locations and formations to be explored by ongoing and future rover missions, including Oxia Planum, which, dated at 4.0 Ga, is the most ancient Mars location targeted for investigation yet.
... This holds particularly true when birds are concerned. As for the fish, A. crassicaudus is commonly found in sediments correlated with all the phases of the Messinian Salinity Crisis in the Mediterranean area (e.g., Carnevale et al. 2019). Sciaenids exhibit a wide stratigraphic and geographic distribution during the Neogene (Bannikov et al. 2018). ...
... Coastal and freshwater fish assemblages in the post-evaporitic Messinian of italy share numerous affinities throughout the Italian Peninsula (Carnevale et al. , 2019. This is particularly evident between Tuscany and Piedmont, even though it could just result from them being better sampled than other sites (i.e., in Emilia Romagna, Marche, and Abruzzo, for example). ...
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The latest Messinian Monticino Quarry fissure fillings, near Brisighella in Italy, are well known for their diverse mammal fauna. Conversely, little is known about other vertebrates from this rich site. beside presenting an overview of the mammals, here we describe fish, amphibian, reptile, and bird remains, identifying three, eight, at least 18, and five taxa, respectively. Some of these represent the oldest occurrences either worldwide (an erycine snake related to Eryx jaculus, Malpolon, and otidids birds) or locally (common toads in the Apennine Peninsula and eremiadine lacertids in Europe, the latter being also the only known occurrence of these lizards in Italy). Altogether, the vertebrate fauna from Monticino Quarry is indicative of a complex palaeoenvironment, which was warm and somehow dry. water bodies were indeed present though, as well as a patchy set of habitats with loose or sandy soils, rocky outcrops, open biotopes, and (even though maybe to a lesser extent) wooded areas. Furthermore, estuarine or lagoonal conditions were likely present near the depositional site. From a palaeobiogeographical point of view, the non-mammal vertebrate assemblage from Monticino Quarry shows considerable Afro-Asian affinities at least for some components. In the context of the Italian late Miocene continental vertebrate assemblages, that of Monticino is remarkably similar to those from the Piedmont basin, especially if compared with the Tusco-Sardinian area, Apulian Platform, Calabrian Arc and Sicily. This is consistent with the palaeobiogeographic scenario defined by mammal assemblages, with northcentral Italy, Tuscany and Sardinia, Apulian Platform, and Calabria and Sicily representing separated bioprovinces.
... Furthermore, several evidences based on fossil occurrence during the Primary Lower Gypsum (PLG) phase (i.e. CN, foraminifers, molecular fossils, diatoms, fishes, and echinoids; Landini and Sorbini, 1989;Lacour and Nèraudeau, 2000;Néraudeau et al., 2002;Carnevale et al., 2019;Pellegrino et al., 2021) argue against a biotic crisis associated with the MSC onset. Although the MSC time interval coincides with the absence of foraminifers (i.e. ...
Article
During the Messinian Salinity Crisis (MSC, 5.97–5.33 Ma) the synergy between tectonic and climatic processes promoted the deposition of evaporites throughout the Mediterranean Basin. The disappearance of calcareous nannofossils and foraminifers at the MSC onset has been traditionally interpreted as the result of the establishment of hypersaline conditions in the water column, which were lethal for most marine eukaryotes. Here, we investigate the calcareous micro and nannofossil assemblages from the Sorbas Basin (SE Spain) spanning the onset and the first phase of the MSC (~ 5.9–5.7 Ma) in order to test whether their disappearance was related to adverse environmental conditions and/or to preferential dissolution of biogenic calcite. Micropaleontological analyses and petrographic observations suggest that the disappearance of calcareous fossils was the result of a taphonomic bias related to paleoceanographic changes that favoured the oxidation of both pyrite and organic matter at the sea floor. A poorly preserved benthic and planktic assemblage and aragonitic laminated sediment characterize the deposits of the first phase of the MSC from 5.93 Ma onward; aragonite deposition was likely bio-mediated in response to enhanced photosynthesis, as suggested by its δ¹³C signature. The fossil assemblage preserved in these deposits indicates a shallowing upward trend of the basin with respect to the pre-evaporitic phase, and the establishment of a lagoon-like environment characterized by episodic improvement of the connections with the open sea. We conclude that the temporary disappearance of calcareous fossils approximately at the MSC onset was related to ongoing basin restriction, which increased the sensitivity of the basin to fresh-water input and temperature changes, promoting marked fluctuations of the redox conditions in bottom sediments. This finding challenges the view of a marine biotic crisis related to hypersaline conditions at the MSC onset and opens new perspectives on the paleoenvironmental conditions that characterized this time interval.
... Thus, along the Pliocene coasts of Tuscany, a largely modern carcharhine assemblage coexisted along with "Miocene survivors" among both the sharks (e.g., the extinct Pachyscylli um distans (Probst, 1879) and Megascyliorhinus mio caenicus (Antunes & Jonet, 1970); Manganelli & Spadini 2019;Collareta et al. 2020b) and the rays (e.g., members of the extant genera Aetobatus Blainville, 1816 and Anoxypristis White & Moy- Thomas, 1941;Collareta et al. 2017Collareta et al. , 2021b) -an observation that evokes a rather gradual transition from the preevaporitic Mediterranean assemblages of the Tortonian and Messinian (e.g., Menesini 1967Menesini , 1969Arambourg 1927;Ward & Bonavia 2001) through the Messinian Salinity Crisis (cf. discussion in Carnevale et al. 2019) and the subsequent Pliocene. ...
Here we report on two fossil teeth attributed to the blacktip shark Carcharhinus limbatus (Elasmobranchii: Carcharhiniformes: Carcharhinidae) from lower Pliocene (ca. 5.1-4.5 Ma) marine deposits of Tuscany (central Italy). A survey of the palaeoichthyological literature and online resources reveals that fossils of C. limbatus have been reported from Neogene and Quaternary deposits of the Americas and Indo-Pacific Asia. Therefore, the Tuscan specimens described herein represent the first occurrence of C. limbatus as a fossil from both Europe and the Mediterranean Basin. Our finds suggest that the blacktip shark reached a circumglobal distribution in warm waters (including the then warmer-than-today Mediterranean Sea) not later than the Pliocene; at the same time, they demonstrate that the continued study of the relatively well-investigated Neogene elasmobranch faunas of Italy can still result in significant novelties. Further investigations on other shark tooth assemblages from shallow-marine, warm-water, nearshore deposits are likely to reveal new occurrences of C. limbatus in the Pliocene of the Mediterranean Sea and elsewhere (e.g., along the palaeontologically under-explored Atlantic and Pacific coasts of Africa).
... In the photic marginal settings, this stage is recorded by up to 16 depositional cycles given by an alternation of primary shallow-water evaporites and dark organic-rich euxinic shales (PLG unit;Lugli et al., 2010). This succession is mostly barren of fossils, an exception being fish (Carnevale et al., 2019) and some marine mollusks found in the shale hemicycles of the Sorbas basin (Betic Cordillera, Southern Spain; Roveri et al., 2020;Saint-Martin et al., 2000). In deeper poorly oxygenated depositional settings, this interval is recorded by organic-and dolomite-rich evaporitic-free deposits, barren of foraminifera (FBI; Manzi et al., 2018) and containing only rare high-salinity tolerant organisms (nannofossils and pteropods; see Manzi et al., 2007). ...
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We present a new ⁸⁷Sr/⁸⁶Sr curve for the time interval between 6.5 and 5.5 Ma of the Ain El Beida section (AEB, Atlantic side of Morocco). The location and the precessional‐scale resolution of our curve allow to investigate the possible relationships between global paleoclimatic and palaeoceanographic trends and the high‐amplitude hydrological changes of the Mediterranean during the Messinian salinity crisis (MSC). The values, obtained from planktic foraminifers, plot in the upper portion of the global ocean curve and show long‐ (at eccentricity scale) and short‐term (precessional scale) oscillations; the latter are more evident during the maximum of eccentricity for both the 100 and 400 ka components, between 6.00 and 5.85 Ma. Negative ⁸⁷Sr/⁸⁶Sr anomalies are observed at insolation minima in phase with δ¹⁸O maxima, marking the glacial stages from TG34 to TG26. It is unclear whether these oscillations originated from local (river runoff) or global (glacial/interglacial) climatic forcing. Conversely, the evaporites of the coeval Primary Lower Gypsum (5.97–5.62 Ma) accumulated in the Mediterranean at insolation minima show positive anomalies at glacial stages TG32, 30, 28, and 26. Such an opposite trend is possibly related to the greater sensitivity of the Mediterranean Sea to river runoff during insolation maxima compared to the global ocean, as recorded at the Moroccan Atlantic margin. These observations and the deviation toward lower values of the Mediterranean Sr isotope curve starting at around 6.5 Ma, support the hypothesis of a progressive restriction of the Atlantic connections since the early Messinian, leading to the MSC.
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Gypsum makes up about one fifth of giant salt deposits formed by evaporation of seawater throughout Earth’s history. Although thermodynamic calculations and precipitation experiments predict that gypsum precipitates when the salinity of evaporating seawater attains about 110 g kg⁻¹, gypsum deposits of the Mediterranean Salt Giant often bear the geochemical signature of precipitation from less saline water masses. Addressing this geochemical riddle is important because marine gypsum deposition and continental gypsum erosion affect the global carbon cycle. We investigated gypsum deposits formed in the marginal basins of the Mediterranean Sea during the Messinian Salinity Crisis (about 6 million years ago). These often bear low-salinity fluid inclusions and isotopically light crystallization water, confirming previous published reports that the Mediterranean Salt Giant harbors low-salinity gypsum deposits. A geochemical model constrained by fluid inclusion salinity and isotope (⁸⁷Sr/⁸⁶Sr, δ³⁴SSO4, δ¹⁸OH2O, δDH2O) measurements excludes that Ca²⁺- and SO4²⁻-enriched continental runoff alone provides the trigger for gypsum precipitation at low salinity. We propose that, concurrent with the prevalent evaporative conditions and with Ca²⁺- and SO4²⁻-bearing runoff, the biogeochemical sulfur cycle is capable of producing a spatially-restricted and temporally-transient increase of Ca²⁺ and SO4²⁻ within benthic microbial mats, creating local chemical conditions conductive to gypsum precipitation. This hypothesis is supported by the presence of dense packages of fossils of colorless sulfur bacteria within gypsum in several Mediterranean marginal basins, together with independent geochemical and petrographic evidence for an active biogeochemical sulfur cycle in the same basins. Should this scenario be confirmed, it would expand the range of environments that promote marine gypsum deposition; it would also imply that an additional, biological coupling between the calcium, sulfur and carbon cycles exists.
Article
Primary gypsum represents an excellent paleobiological archive due to its early and fast growth, favoring the preservation of delicate biomineralized structures. The Mediterranean region is renowned for evaporite deposits that formed during the Messinian salinity crisis (MSC), an event that supposedly annihilated most of the marine biota. However, the Messinian evaporites have been scarcely studied for their fossil content. Abundant nano-sized planktic diatoms and associated organic matter are observed for the first time in bottom-grown gyp-sum crystals that formed during the early stage of the MSC in different marginal basins of the western Mediterranean. This discovery increases our knowledge of the Messinian biota and reveals that nano-sized planktic diatoms played a prominent role in carbon and silicon export during gypsum deposition. The co-occurrence of these diatoms with larger diatoms, possibly associated with a deep chlorophyll maximum, suggests that Messinian gypsum formed in stratified and relatively deep basins (far below the photic zone), typified by marine conditions in the upper water column. The nano-sized planktic diatoms may have taken advantage of the hydrological reconfigurations experienced by the Mediterranean since the onset of the MSC. This study confirms that primary gypsum represents a promising archive of information for elucidating the marine biotic response to an ancient environmental crisis.
We report on new fossils of chitinophosphatic brachiopods from Upper Pliocene (Piacenzian) marginal-marine clayey sands cropping out at La Serra (Tuscany, central Italy). Based on their morphological characters, these specimens are here assigned to the linguliform family Lingulidae and tentatively recognised as belonging to the extant genus Lingula. This is the first record of lingulides from the Mediterranean Pliocene, as well as the geologically youngest one from the Mediterranean Basin. Since the family Lingulidae has previously been reported from the upper Miocene of Italy, the occurrence of lingulides in the Pliocene of Tuscany evokes the persistence of lingulide brachiopods within the Mediterranean Sea during the Messinian Salinity Crisis or their recolonization of this basin shortly after the Miocene-to-Pliocene transition. The eventual extirpation of the family Lingulidae from the Mediterranean Sea could have resulted from the progressive cooling of seawater and loss of shallow-water coastal environments that occurred in Pleistocene times.
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Abstract: We report about the discovery of a mysticete cetacean neurocranium located inside a block of oolitic limestone, corresponding to the Terminal Carbonate Complex (TCC) of the island of Mallorca. This unit is directly related to the Messinian Salinity Crisis (MSC) that occurred in the Mediterranean 5.97-5.33 Ma ago. The importance of the finding lies mainly in that it is the first and only record, so far known, of a fossil whale in the Messinian through the Mediterranean, since it is assumed that as a consequence of extreme environmental conditions during MSC, the great cetaceans went temporarily extinct from the Mediterranean. The record of the MSC in the Balearic Islands is nearer and consistent to the models that suggest two evaporitic episodes, separated by a brief restitution of sea-level would have occurred (5.77-5.60 Ma) likely provoked the deposition of the TCC in the marginal zones, thus possibly allowing the sporadic entrance of cetacean in the Messinian Mediterranean. Key words: Mysticeti, Messinian Salinity Crisis, Terminal Carbonate Complex, Mallorca, Western Mediterranean.
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The fossil record of sciaenid fishes (based on both otoliths and osteological finds) from the Neoge-ne of the Paratethys is reviewed. The species Labrax (=Morone) multipinnatus Gorjanović-Kramberger, 1882 from the Sarmatian of Croatia (Sv. Nedelja) that was originally described as a sea bass of the family Moronidae is reassigned to the extant croaker genus Argyrosomus. Two new genera and species of sciaenid fishes are described based on a single skeleton each, namely Landinisciaena popovi gen. et sp. nov. from the Tarkhanian (Lower/Middle Miocene) North Shir-vanskaya Formation exposed along the Pshekha River, North Caucasus, Russia, and Croatosciaena krambergeri gen. et sp. nov. from the Sarmatian s.s. (Middle Miocene) deposits of Dolje in the nearby of the city of Zagreb, Croatia. The holotype of the former taxon possesses a well-preserved saccular otolith in situ (first record of in situ croaker otolith in Europe) similar to the otoliths of the extant genus Atractoscion. The otolith-based Ottnangian species Atractoscion elongatissimus Schwarzhans, 1993 is transferred to the genus Landinisciaena gen. nov. A comprehensive revision of the Neogene otholith-based sciaenid record from Paratethys resulted in the recognition of three new genera (Chaoia gen. nov., Pontosciaena gen. nov. and Leptosciaena gen. nov.) as well as in the description of a new species L. caputoi sp. nov. from the uppermost Messinian (Lago-mare phase) of Italy. A number of known otholith-based species were either synonymized or assigned to a new generic affiliation. The complex biogeographic history of the Neogene Paratethyan sciaenids (based on both otoliths and skeletal records) is discussed, resulting in the recognition of a vanished "sciaenid bioprovince".
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In modern stromatolites, mineralization results from a complex interplay between microbial metabolisms, the organic matrix, and environmental parameters. Here, we combined biogeochemical, mineralogical, and microscopic analyses with measurements of metabolic activity to characterize the mineralization processes and products in an emergent (<18 months) hypersaline microbial mat. While the nucleation of Mg silicates is ubiquitous in the mat, the initial formation of a Ca‐Mg carbonate lamina depends on (i) the creation of a high‐pH interface combined with a major change in properties of the exopolymeric substances at the interface of the oxygenic and anoxygenic photoautotrophic layers and (ii) the synergy between two major players of sulfur cycle, purple sulfur bacteria, and sulfate‐reducing bacteria. The repetition of this process over time combined with upward growth of the mat is a possible pathway leading to the formation of a stromatolite.
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In marginal Mediterranean sub-basins, the early phase of the Messinian salinity crisis (MSC) is recorded by cyclic successions of gypsum and shales, which in deeper parts of the sub-basins make lateral transition into organic-rich shales, marls, and carbonates. The cyclic stacking pattern of the gypsum-bearing sequences is assumed to reflect periodic paleoenvironmental change induced by precession-driven climate perturbations, with the assumption that shales reflect humid climate (precession minima) while gypsum reflects arid conditions (precession maxima). However, this correlation has not been verified to date, mostly because of the scarcity of microfossils, the most commonly used tools for the reconstruction of precession-driven paleoenvironmental change. Such change can, instead, be reconstructed through the study of the deeper water counterparts of gypsum (marly and carbonate layers) with geochemical indicators (major and trace elements, molecular fossils), which provide insight on climate and aquatic productivity. We used this approach to study a section from the Piedmont Basin (NW Italy) where the onset of the MSC is archived in a sequence of organic-rich sediments. This sequence displays distinct lithological cyclicity, evidenced by the repetition of couplets of organic-rich shales and marls, either bioturbated (in the pre-MSC part of the section) or laminated (during the MSC). The influence of orbitally-driven (precession) climate oscillations is demonstrated by fluctuations of Ti/Al, Si/Al, Mg/Al, K/Al, Zr/Al, and Ba/Al ratios that are in phase with lithological cyclicity. These fluctuations are interpreted to reflect alternation of humid (shales, deposited during precession minima) and arid (bioturbated and laminated marls, deposited during precession maxima) phases, dominated by fluvial and aeolian transport of detrital material, respectively. The cyclicity of the element ratios is mirrored by changes in organic carbon content and molecular fossil inventory. In particular, the distribution of long-chain n-alkanes and their degree of preservation reveal that humid phases at times of precession minima were typified by the maximum input of degraded terrestrial organic matter driven by enhanced riverine runoff, which promoted water column stratification. Coeval increase in Ba content, a common paleoproductivity proxy, agrees with enhanced nutrient supply during humid periods, promoting phases of eutrophication in the basin. Lithological and geochemical changes are observed in MSC sediments deposited at times of precession maxima, evidenced by the replacement of pre-MSC bioturbated marls by laminated marls rich in filamentous fossils corresponding to the remains of probable colorless sulfide-oxidizing bacteria. Such changes reflect an intensification of water column stratification after the onset of the MSC, possibly related to the combined effect of persistent freshwater inflow and basin isolation, preluding the advent of gypsum precipitation.
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New palaeontologic, sedimentologic, and Sr isotope data allow to reconstruct a high‐resolution chronostratigraphic and palaeoenvironmental framework of the continentalmarine transition at the Miocene–Pliocene boundary in the Sorbas Basin (Betic Cordillera, South‐eastern Spain). The presence of Reticulofenestra zancleana, Ceratolithus acutus, and R. pseudoumbilicus in a marine horizon sharply overlying the continental deposits of the Zorreras Member indicates that the Messinian salinity crisis ended in the Sorbas Basin synchronously with the other Mediterranean basins at the base of the Zanclean, within the MNN12a biozone. Our results suggest that the Zanclean flooding turned the Sorbas Basin into a shallow bay with limited exchange with the main Mediterranean basin, probably through narrow seaways connecting also the Almeria, Nijar, and Vera basins. Our results do not confirm previous hypotheses envisaging an older age for the return to fully marine conditions in the Sorbas Basin and is in a good agreement with the reconstructions suggesting that the Zorreras Mb. continental deposits are the local time‐equivalent of the latest Messinian Lago‐Mare phase during the last stage of the salinity crisis.