# Unravelling land-sea correlations during the Pliocene-Pleistocene in the Mediterranean area

Adele BERTINI*, Maria MARINO, Nathalie COMBOURIEU-NEBOUT, Angela GIRONE, Patrizia MAIORANO, Gabriele NICCOLINI & Jean-Pierre SUC

A. Bertini, Dipartimento di Scienze della Terra, Università degli Studi di Firenze, Via G. La Pira 4, I-50121 Firenze, Italy; adele.bertini@unifi.it
*corresponding author

M. Marino, Dipartimento di Scienze della Terra e Geoambientali, Università degli Studi di Bari Aldo Moro, Via E. Orabona 4, I-70125 Bari, Italy; maria.marino@uniba.it

N. Combourieu-Nebout, HNHP, Histoire Naturelle de l'Homme Préhistorique, UMR 7194 - CNRS/MNHN, 1 rue René Panhard, F-75013 Paris, France; nathalie.nebout-combourieu@mnhn.fr

A. Girone, Dipartimento di Scienze della Terra e Geoambientali, Università degli Studi di Bari Aldo Moro, Via E. Orabona 4, I-70125, Bari, Italy; angela.girone@uniba.it

P. Maiorano, Dipartimento di Scienze della Terra e Geoambientali, Università degli Studi di Bari Aldo Moro, Via E. Orabona 4, I-70125 Bari, Italy; patrizia.maiorano@uniba.it

G. Niccolini, Dipartimento di Scienze della Terra, Università degli Studi di Firenze, Via G. La Pira 4, I-50121 Firenze, Italy; gabriele.niccolini@unifi.it
J.-P. Suc, Sorbonne Université, CNRS-INSU, Institut des Sciences de la Terre Paris, ISTE, UMR 7193, F-75005 Paris, France; jeanpierre.suc@gmail.com

KEY WORDS - Pollen, Flora, Vegetation, Calcareous plankton, Palaeoenvironments, Quaternary.

**ABSTRACT** - In this paper, we synthetised pollen, foraminiferal, and calcareous nannofossil data from marine and continental successions in Italy to investigate the complex dynamics of climate-induced shifts in terrestrial and marine ecosystems during the Late Pliocene and Early to Middle Pleistocene. Additionally, by focusing on three key case studies derived from interdisciplinary research of marine sedimentary successions (Semaforo, Vrica-Santa Lucia and Montalbano Jonico) we present concurrent evidence from both marine and continental proxies, offering a refined perspective on the environmental history of the Mediterranean. This integrated approach enhances the precision of reconstructing climate system interactions during critical intervals of the Quaternary, mitigating potential discrepancies arising from independent chronologies and methods of study. Overall, the synthesis and case studies contribute to a more comprehensive understanding of the role of the Mediterranean in global climate variability, emphasising the sensitivity of the region to both continental and marine climatic drivers throughout the Quaternary.

## INTRODUCTION

The Mediterranean region has long been recognised as a key area for investigating climate variability and environmental change, particularly from approximately 3 million years (Ma) onwards (e.g., Colleoni et al., 2015; De Boer et al., 2021; Berner et al., 2022; Mondanaro et al., 2024). This interval marks the onset of large-scale climatic oscillations driven by glacial-interglacial (G/I) cycles that shaped global climate system. The unique geographical setting of the Mediterranean, situated at the crossroads of temperate, subtropical, and arid climatic zones, makes it particularly sensitive to both regional and global climatic shifts. This sensitivity, combined with the semi-enclosed nature of the Mediterranean Basin, provides an exceptional archive for reconstructing climate dynamics and biotic responses during key phases at the Pliocene-Pleistocene transition (PPt; e.g., Petrick et al., 2018; McClymont et al., 2023) and the Early-Middle Pleistocene Transition (EMPT; Head & Gibbard, 2015). The complex interplay of G/I cycles, sea-level changes, and orbital forcing in the Mediterranean region has left behind high-resolution records in both marine and terrestrial sedimentary archives. These records capture critical responses to global cooling, changes in insolation, and variations in atmospheric and oceanic systems, offering valuable insights into the processes driving

environmental and ecological transformations. Notably, marine proxies such as foraminifer and calcareous nannofossil assemblages provide reconstructions of sea-surface features (temperature, salinity, turbidity, nutrient content, stratification/mixing) (Hemleben et al., 1989; Beaufort et al., 1999; Bárcena et al., 2001; Bauman et al., 2005), while pollen records deliver an unparalleled view of land flora composition, vegetation dynamics and regional climate conditions (Suc et al., 2018; Bertini & Combourieu-Nebout, 2023 and references therein). By integrating terrestrial and marine proxy records, it is possible to trace the timing, magnitude, and mechanisms of environmental changes. This approach enables a deeper understanding of how ecosystems responded to major climatic drivers. Numerous studies have used these proxies to highlight the Mediterranean's climatic fluctuations over orbital and millennial timescales (e.g., Martrat et al., 2004; Fletcher & Sánchez-Goñi, 2008; Suc et al., 2010, 2018; Ausín et al., 2015; Bertini et al., 2015; Combourieu-Nebout et al., 2015; Rohling et al., 2015; Maiorano et al., 2016a, b; Tzedakis et al., 2017; Marino et al., 2020; Toti et al., 2020), though some have been limited by lower-resolution datasets (see references in Bertini, 2010; Bertini et al., 2010; Magri et al., 2017; Bertini & Combourieu-Nebout, 2023). Nevertheless, all previous contributions have significantly advanced our comprehension of environmental shifts in the region.
The Mediterranean's role in deciphering Quaternary climate dynamics extends beyond regional relevance; it has broader implications for understanding global climate systems. The challenge of correlating terrestrial and marine records is magnified by the region's tectonic activity, sea-level fluctuations, and hydrological variability, all of which add layers of complexity to palaeoenvironmental reconstructions (e.g., Nomade et al., 2019; Marino et al., 2020; Dumitru et al., 2021; Lionello et al., 2023). However, the integration of these records (combining palynological data with marine micropalaeontological evidence) offers a robust framework for unravelling the intricate land-sea correlations during the Quaternary. Understanding these complex interactions is not only essential for reconstructing past environmental conditions, but also for predicting future responses to ongoing climate change in this highly vulnerable region.

In this contribution, we critically synthesised key information to reconstruct Mediterranean environments during the Pliocene and Pleistocene, by integrating data from both continental and marine sedimentary successions which were calibrated using independent time-diagnostic tools. Additionally, we aimed to emphasise, through specific case studies, the critical role that interdisciplinary studies - combining palynological data, calcareous plankton, and other biogeochemical proxies from marine successions - play in constructing comprehensive climate change assessments. Indeed, by comparing and correlating data from both terrestrial and marine contexts, we obtained insights into synchronous climatic and oceanographic-atmospheric events. This integrative approach not only highlights the intrinsic connections between these two seemingly distinct realms but also offers a more nuanced understanding of how global climate dynamics is expressed across different ecosystems concurrently.

## MATERIALS AND METHODS

This study adopted a multidisciplinary approach to evaluate the effectiveness of land-sea correlations, synthesising bibliographic data from key reference sites throughout Italy (Figs 1 and 2). For each of the first three stages of the Quaternary (Gelasian, Calabrian, Chibanian) we reconstructed terrestrial environments using pollen data to analyse floristic composition and changes in vegetation. Concurrently, marine conditions are assessed through records of planktonic foraminifera and calcareous nannofossils, which provided insights into sea surface temperature, productivity, and ocean circulation patterns. When working with different natural archives, such as lacustrine versus marine deposits, we selected those that have a robust chronostratigraphic and geochronologic framework thanks to the use of temporally diagnostic tools, such as radiometric dating, magnetostratigraphy, and biostratigraphy.

A notable advantage arises when both terrestrial and marine proxies are derived from the same marine archive, as this facilitates direct correlation and enhances the precision of palaeoenvironmental reconstructions. This advantage is particularly significant due to the remarkable ability of pollen grains to be deposited and preserved within various environmental contexts, including

marine settings. Pollen grains become incorporated into sediments along with marine plankton, contributing to the accumulation of deposits in marine basins. This integrated approach enabled us to capture the differing response times of terrestrial and marine systems to substantial climatic shifts in the Mediterranean region. By aligning terrestrial pollen data with marine planktonic microfossils, we can effectively bridge gaps that often arise from the independent application of chronostratigraphic and geochronologic schemes, thus resolving issues of temporal misalignment. The robustness of marine archives in this context is crucial, as they allow for the simultaneous recording of land-sea interactions, facilitating comprehensive testing and validation of these correlations. This, in turn, enhances our understanding of environmental changes. By focusing on three key case studies, including Semaforo, Vrica-Santa Lucia, and Montalbano Jonico sites, we illustrated how interdisciplinary approaches to marine successions are indispensable for detecting synchronous climatic and environmental shifts across the Mediterranean region (Figs 3-6). Furthermore, following the two preceding insights into the Italian sites, comparisons were extended to coeval continental and/or marine sites within the Mediterranean region, as well as to key sites at a broader extra-Mediterranean scale. In Fig. 2 the Astronomically Tuned Neogene Time Scale (ATNTS2004) by Lourens et al. (2004) is used as reference, integrating marine sedimentary cycles, palaeomagnetic reversals, and biostratigraphy from Mediterranean deep-sea cores (Ocean Drilling Program-ODP sites). This chronology provides a robust framework based on oxygen isotope, Milankovitch cyclicity, sapropel layers, and calcareous plankton bioevents. Sapropel stratigraphy aligns with the syntheses by Lourens et al. (1996a, 2004). The Pliocene-Pleistocene chronostratigraphy follows the International Union of Geological Sciences (IUGS) standard (June 2009; Head et al., 2008; Walker & Geissman, 2009). Additional chronostratigraphic comparison is made with Gradstein et al. (2004) and Gradstein & Ogg (2020) for clarity. Biozones of calcareous nannofossils are based on Rio et al. (1990), and planktonic foraminiferal zones on Cita (1975, amended) and Lirer et al. (2019). Astronomical ages for biozone boundaries follow Lourens et al. (2004) and Raffi et al. (2006). The pollen records from selected Italian continental and marine sites are constrained by palaeomagnetic, marine stratigraphic, and biochronological markers (e.g., mammalian biozones) (Fig. 2). Stratigraphic and geological data for each site, along with references for detailed pollen data, are from Bertini (2010), Bertini et al. (2010), Sadori et al. (2013), Combourieu-Nebout et al. (2015), Magri et al. (2017), Bertini & Combourieu-Nebout (2023), along with relevant references therein.

## FROM LATEST PIACENZIAN TO CHIBANIAN: A COMPREHENSIVE SYNTHESIS OF KEY ENVIRONMENTAL CHANGES IN ITALY

This overview focuses on the first three stages of the Pleistocene, emphasising significant climate changes, including the strengthening of the Northern Hemisphere Glaciation (NHG) around 2.7 Ma (e.g., Ravelo et al.,
2004; Bartoli et al., 2005, 2011; Mudelsee & Raymo, 2005; Ruggieri et al., 2009; Kabot-Bhar et al., 2021; McClymont et al., 2023) following the mid-Piacenzian Warm Period (mPWP, 3.3-3 Ma), characterised by reduced global ice volume and elevated pCO₂ (e.g., de la Vega et al., 2020). It also analyses the EMPT (1.25-0.4 Ma; Head & Gibbard, 2015), which resulted in intensified glacial phases in the Mediterranean region and at global scale, leading to increased aridity on land and more pronounced fluctuations in marine environments.

### Latest Piacenzian-Gelasian (Figs 1-4)

Since the late Piacenzian, the shift from relatively warm to cooler climates paved the way for the high-magnitude G/I cycles of the Quaternary. The Gelasian (~2.58-1.8 Ma, Marine Isotope Stages, MIS 103-MIS 65/64), marking the beginning of this period, is characterised by the onset of large-scale NHG driven by obliquity (41 kyr). This transition profoundly impacted global climate patterns, leading to significant changes in both terrestrial and marine environments throughout the Mediterranean region.

**MARINE RECORDS** - The Pliocene-Pleistocene transition marks the onset of the profound G/I fluctuations with sawtooth pattern of benthic $δ^{18}$O, which were the main feature of Pleistocene Epoch as recorded by multiple data (e.g., Shackleton et al., 1995; Lisiecki & Raymo, 2005). The transition is well documented in the Mediterranean marine archives thanks to well exposed on land sedimentary successions, which allowed high resolution

cyclostratigraphy and biostratigraphy starting from the Piacenzian (e.g., Zachariasse et al., 1989; Sprovieri et al., 2006). The $δ^{18}$O records, both at low (e.g., Lourens et al., 1992; Howell et al., 1998; Kroon et al., 1998; Serrano, 2020) and high resolution (Becker et al., 2005; Grant et al., 2022; Addante et al., 2024a-b, 2025; Zanola et al., 2024), evidence the high latitude climate variability but also the influence of low latitude African monsoon activity (Rohling et al., 2015), which determined the deposition of sapropel layers in marine sediments at the precession scale (Hilgen, 1991; Rohling et al., 2015). These climate conditions affected the calcareous plankton evolution in response to progressive lower sea surface temperature, variation of salinity and nutrient availability, and structure of photic waters (stratification/mixing, nutricline/thermocline depth). Related to the cooling trend through PPt and the NHG and maybe to shallowing nutricline and mixing (Schueth & Bralower, 2015; Tangunan et al., 2018) is the prominent drop of warm and oligotrophic nannofossil *Discoaster* spp. documented in many deep sea cores recovered during several ODP expeditions (e.g., ODP Legs 13, 42, 107, 160, 161; see http://www-odp.tamu.edu/publications/) and on land marine sections, such as Monte San Nicola (Driever, 1988; Channell et al., 1992; Rio et al., 1998; Addante et al., 2025), Mandorlo (Capraro et al., 2022; Zanola et al., 2024), and Punta Piccola (Castradori et al., 1998) in Sicily, Singa in Calabria (Lourens et al., 1992), along with Marecchia Valley in northern Apennines (Rio et al., 1996). They also enriched the Mediterranean calcareous plankton biostratigraphic

<figure><img src="image_1.png" /><figcaption>Fig. 1 - (color online) Location map of selected Pleistocene marine and continental sites (1-27). 1: Piánico-Sëllere; 2: Leffe; 3: Venice; 4: Cava RDB; 5: Castell'Arquato, Stirone; 6: Lamone, Santerno; 7: Val Marecchia; 8: Lower Valdarno; 9: Upper Valdarno; 10: Colfiorito; 11: Tiberino Basin; 12: F. Ranuccio-Costa S. Giacomo; 13: Isernia; 14: Monticchio; 15: Acerno; 16: Vallo di Diano; 17: Sant'Arcangelo; 18: Montalbano Jonico; 19: Crotone Basin; 20: ODP site 975; 21: Singa; 22: Bianco; 23: Punta Piccola-Capo Rossello; 24: Monte San Nicola; 25: ODP site 976; 26: core KC01B; 27: ODP site 964. For additional information on sites see the references in Bertini et al. (2010) and Bertini & Combourieu-Nebout (2023).</figcaption></figure>
schemes (e.g., Cita, 1975; Rio et al., 1990; Lirer et al., 2019; see also synthesis in Capraro & Maiorano, 2023), the sapropel stratigraphy and the Astronomical Time Scale (Lourens et al., 1996a). In the Monte San Nicola section, where the Global Stratigraphic Section and Point (GSSP) of the Gelasian Stage was defined (Rio et al., 1998), the last occurrence of *Discoaster pentaradiatus* (Tan, 1927) has been recently assigned to an age of 2.52 Ma (Addante et al., 2024a), within MIS 100, together with a relevant abundance drop of the genus. Moreover, a first significant occurrence of the polar-subpolar foraminifer *Neogloboquadrina atlantica* (Berggren, 1972) has been recognised during MIS 104, just below the Gelasian GSSP, and in MIS 100. The influxes of this taxon in the Mediterranean Sea were previously reported through the Pliocene-Pleistocene transition (MIS 104 and MIS 100) by other authors (e.g., Lourens et al., 1996a; Caruso, 2004; Becker et al., 2005; Addante et al., 2024b). These calcareous plankton events are a clear signal of climate deterioration and cooling of sea surface temperature (SST), which also emphasise the connection between high and middle latitude climate during the NHG. The southward migration of the Subarctic Front at these times agrees with lower alkenone-SSTs at the Monte San Nicola section (Herbert et al., 2018; Addante et al., 2024a) and is contemporaneous with the occurrences of ice rafted debris (Naafs et al., 2013), lower sea surface temperature (Naafs et al., 2020) and *N. atlantica* (Bolton et al., 2010) at the mid-latitude North Atlantic Integrated Ocean Drilling Program-IODP site U1313.

**TERRESTRIAL RECORDS** - Pollen data indicate a marked transition toward cooler and drier conditions. The prevalent expansion of open steppe-like vegetation and a reduction in forested areas suggest a response to the advancing glaciations. Notably, the palynological records show a decline in thermophilous arboreal taxa particularly those currently distributed in subtropical regions, and an increase in herbaceous plants, reflecting the colder, more arid climate (Bertini, 2010; Suc et al., 2018; Bertini & Combourieu-Nebout, 2023 and references therein).

After the glacial marine isotope stage M2 (~3.3 Ma), the mPWP marked a final global warm phase, characterised by a rise in sea surface temperatures of approximately 3 °C and atmospheric CO2 levels reaching around 400 ppm, comparable to modern anthropogenic levels (e.g., Haywood & Valdes, 2004; Pagani et al., 2009; Bartoli et al., 2011). In central Italy, at Santa Barbara (Upper Valdarno), a rich subtropical-to-warm-temperate forest thrived during this warm phase (~3.0 Ma), following the cooling episodes of M2 and MG2. Dominated by *Taxodium/Glyptostrobus*-type, *Nyssa*, and other thermophilous taxa, this forest gradually transitioned to cooler species such as *Picea* and *Fagus* around 2.8

Ma, indicating a shift towards cooler conditions (Bertini, 2010, 2013). This transition aligns with isotopic evidence of intensifying glaciation and the subsequent reduction or disappearance of thermophilous taxa across central and northern Italy (Bertini, 2010 and references therein).

In southern Italy, Piacenzian pollen records, though more limited, reveal a cool, humid climate with reduced seasonality around 3.31 Ma, especially at Bianco, where forests were dominated by *Picea* and *Abies*, and Mediterranean evergreen taxa experienced a marked decline (Bertoldi et al., 1989). Between 3 and 2.92 Ma, at Punta Piccola (Sicily), high-resolution records suggest alternating warm and arid phases linked to precession cycles, with humid conditions prevalent during the deposition of laminated sediments (Combourieu-Nebout et al., 2004). By 2.5 Ma, increasingly drier conditions and the spread of Mediterranean vegetation were documented at Capo Rossello (Sicily), reflecting a progressive shift towards arid climates during the warm season (Bertoldi et al., 1989; Rio et al., 1990). Since ~3.0 Ma, the intensification of Mediterranean outflow and the onset of NHG further triggered cooler, more seasonal, and drier conditions across the Mediterranean region (Lourens et al., 1996b, 2004). This is reflected in the 41 kyr obliquity-driven G/I cycles observed in Italian pollen records, characterised by alternating forest and steppe vegetation (Bertini, 2010; Combourieu-Nebout et al., 2015). A prominent example is the Semaforo section (Crotone, Calabria) (Fig. 4a-b and c1), where pollen and oxygen isotope data provide evidence of these climatic shifts from 2.46 Ma onwards (Combourieu-Nebout & Vergnaud-Grazzini, 1991; Combourieu-Nebout, 1993, 1995). However, pollen data from other southern Italian sites, such as the Gelasian sites at Le Castella and Gela (Monte San Nicola), reveal a contrasting pattern, where forests expanded during glacials, and open vegetation dominated during interglacials, according to Bertoldi et al. (1989). More recent high-resolution studies at Monte San Nicola refine the previously described pattern, contrasting with the more typical one observed along the Mediterranean littoral (Bertini et al., 2024). Glacial and interglacial intervals are distinctly characterised by the prevalence of steppe taxa and thermophilous arboreal plants, respectively. In northern Italy, G/I cycles at sites like Castell'Arquato and Stirone (Emilia-Romagna) are characterised by subtropical-to-warm-temperate forests, contrasting with coniferous forests, with limited steppe vegetation (Bertini, 2001). In the Upper Valdarno Basin, a significant expansion of herbaceous taxa, including *Artemisia*, was recorded in aeolian deposits during the first G/I cycles (Bertini, 1994, 2010). The Piacenzian-Gelasian transition in the Tiberino Basin shows vegetation responding to G/I cycles through the competition between

Fig. 2 - (color online) Marine and continental fossil correlation from late Piacenzian to Chibanian in Italy and the Mediterranean Sea. On the left, the Standard Biochronostratigraphy. The column Series reports the Pliocene-Pleistocene chronostratigraphy according to the IUGS statement (June 2009), Head et al. (2008), and Walker & Geissman (2009). Planktonic foraminifera biozones are from Cita (1975), emended, and Lirer et al. (2019). Calcareous nanofossil biozones are from Rio et al. (1990). In the column Stages, 1, 2, and 3 correspond to the Greenlandian, Northgrippian and Meghalayan, according to Walker et al. (2018). The Mediterranean δ¹⁸O planktonic foraminifer *Globigerinoides ruber* stacked record is from Lourens (2004, and references therein). Sapropel age/stratigraphy is in agreement with the synthesis in Lourens et al. (1996b, 1998) and Lourens (2004). Land Mammal Ages are from Gliozzi et al. (1997), modified; Faunal Units are from Palombo (2004, 2007), Rook & Martinez-Navarro (2010). For additional information on Italian and Mediterranean sites see the references in Bertini et al. (2010) and Bertini & Combourieu-Nebout (2023).
<figure><img src="image_2.png" /><figcaption>Fig. 3 - (color online) a) Panoramic view of the Crotone area. b) Semaforo section with indication of sapropel layer. c) View on the five parts of the Crotone series. d) Details of the Vrica C section with indications of some sapropel layers. e-f) Details of the Vrica B section with indication of the ash layer and some sapropel layers.</figcaption></figure>

coniferous and thermophilous forests (Pontini, 1997; Martinetto et al., 2014). In northern Italy, the progressive decline of “Taxodiaceae” forests (including Taxodioideae and Sequoioideae) and the rise of *Cathaya* are noted from around 2.7 Ma (Stirone), while in southern Italy, “Taxodiaceae” forests at Semaforo decline from 2.38 Ma, with *Cathaya* expanding significantly between 1.92 and 1.74 Ma (Combourieu-Nebout & Vergnaud-Grazzini, 1991; Klotz et al., 2006; Suc et al., 2010).

Climate reconstructions for the last 200 kyrs of the Piacenzian at Stirone (northern Italy) indicate warm and cold phases with temperature and precipitation values similar to those of the following Gelasian (Fauquette & Bertini, 2003). During the Gelasian, warm phases in

northern Italy exhibited high mean annual temperatures (MATs) and precipitation (MAP), comparable to those of the Zanclean, with temperatures around 12-13 °C during glacials and precipitation exceeding modern values by 200-500 mm. In central Italy (Poggio Rosso, Upper Valdarno), climate amplitude reconstructions from a G/I cycle just before the Gelasian/Calabrian boundary show MATs of 12.5-14 °C and MAP of about 800 mm during glacials (with a minimum of 400 mm) and MATs of 15-19 °C and MAP of 750-1200 mm during interglacials (Bertini et al., 2010). In southern Italy, the Semaforo site reveals higher interglacial temperatures (by at least 2.8 °C annually and 2.2 °C in winter) and 500 mm more precipitation compared to today, during the
period from 2.46 to 2.11 Ma (Klotz et al., 2006). Glacial conditions showed temperatures similar to present, with precipitation remaining equivalent and occurring all year along, although there was a general cooling trend over consecutive G/I cycles.

### Calabrian (Figs 1-6)

As previously noted, the EMPT marked a major global climatic reorganisation that profoundly influenced oceanic and atmospheric circulation, ice sheets, and biota, including early humans (Head & Gibbard, 2015). Lasting until about 0.4 Ma, it saw a shift from 41 kyr obliquity-driven cycles of the Early Pleistocene to the 100 kyr G/I cycles of the later Pleistocene. Several severe glaciations occurred, such as MIS 36, 24, and 22, particularly impacting the Northern Hemisphere.

### MARINE RECORDS
The Calabrian Stage, covering MIS 65/64 to MIS 19 interval, is well represented in the Mediterranean region due to the on land marine successions well exposed mainly in the Crotone Basin (Calabria) and Sicily, and deep-sea cores (ODP sites 964 and 967, KC01B) (see a synthesis in Cita et al., 2006, 2008, and reference therein). The Vrica section, housing the Calabrian GSSP, consists of spectacularly exposed blue hemipelagic marly clays, astronomically calibrated thanks to the occurrence of several sapropel layers (Lourens et al., 1996b, 1998). It is a continuous succession with high sedimentation rate (Pasini & Colalongo, 1994; Rio et al., 1996; Suc et al., 2010) that made possible wide interdisciplinary study including marine invertebrate (molluscs, ostracods, planktonic and benthic foraminifera) and pollen (Suc et al., 2010 and references therein) as well as oxygen isotope stratigraphy and magnetostratigraphy (Lourens et al., 1996a, 1998). Among the plethora of biostratigraphic events and data, it is worth to mention the signals of climate deterioration close to the Calabrian GSSP that are the extinction of the warm nannofossil *Discoaster brouweri* (Tan, 1927, emend. Bramlette & Riedel, 1954) (the last survivor of the genus decimated during the PPt), the first increase in abundance of polar-subpolar *Neogloboquadrina pachyderma* (left-coiled) (Lourens et al., 1996a), and the first occurrence of the cold guest benthic foraminifer *Hyalinea balthica* (Schröter, 1783), the latter also recorded at the Rossello section in MIS 53 (Caruso, 2004). Other significant biostratigraphic events in the Calabrian Stage are the appearance of medium-sized *Gephyrocapsa* (4-5.5 µm) (*G. oceanica* Kamptner, 1943; *G. caribbeanica* Boudreaux & Hay in Hay et al., 1967), the last occurrence of *Calcidiscus macintyrei* (Bukry & Bramlette, 1969), the first occurrence of large *Gephyrocapsa* (> 5 µm), and the last occurrence of *Helicosphaera sellii* (Bukry & Bramlette, 1969) (e.g., Rio et al., 1990). Following a temporary disappearance/rarity of *N. pachyderma* from MIS 43 to MIS 37, a new increase of the species is recorded in the glacial MIS 36, in agreement with its distribution in the Montalbano Jonico section (MJS, Gironé et al., 2013a) and ODP site 967 (Lourens et al., 1998), almost synchronously with data from North Atlantic Ocean (ODP site 607, Ruddiman et al., 1989; see synthesis in Maiorano et al., 2010). The MIS 31, considered as a super interglacial analogous of the present Holocene (Maldonado et al., 2014) and the last significant warm interglacial of the obliquity-

dominated climate regime (Scherer et al., 2008), is a significant wide scale period of climate amelioration. It is recorded in the Mediterranean δ¹⁸O record (Lourens et al., 1996a, 2004; Lourens, 2004) and the MJS, and is accompanied by a biostratigraphic event maybe of climate significance, the lowest common occurrence of the extinct nannofossil *Reticulofenestra asanoi* (Sato & Takayama, 1992) (Maiorano et al., 2004). A severe cold glacial during the Calabrian is represented by MIS 22 as recorded in the MJS. It marks the beginning of higher amplitude G/I oscillations and is characterised by the base of temporary disappearance of the warm nannofossil *Gephyrocapsa omega* (Bukry, 1973) (Maiorano et al., 2004) because of the cooling trend during the first major ice volume-induced sea level decrease of the Pleistocene (Shackleton & Opdyke, 1976).

### TERRESTRIAL RECORDS
In southern Italy, extensive palynological records provide evidence of climatic changes across this interval. Notable sites include Vrica (ca. 2.2-1.36 Ma; e.g., Combourieu-Nebout et al., 1990), Santa Lucia (1.356-1.24 Ma, Joannin et al., 2007), and Valle di Manche (0.87-0.73 Ma, e.g., Capraro et al., 2005, 2022) in the Crotone Basin, along with marine and continental records from the MJS (1.24-0.9 Ma, Joannin et al., 2008; 0.82-0.774 Ma, Maiorano et al., 2008; Bertini et al., 2015; Toti, 2015; Toti & Bertini, 2018; Nomade et al., 2019) and Sant'Arcangelo successions (San Lorenzo cycle: ca. 1.1-0.8 Ma, Sabato et al., 2005) in Basilicata. Pollen assemblages indicate a dynamic interplay between open vegetation and forested landscapes, characterised by steppe taxa (e.g., *Artemisia*, *Ephedra*) and warm temperate forest species (e.g., *Quercus*, *Carya*). Contrastingly, northern Italy's records (e.g., Lamone: Fusco, 2007; Leffe: Ravazzi & Rossignol Strick, 1995; Muttoni et al., 2007; Stirone: Lona & Bertoldi, 1972; Bertolani Marchetti et al., 1979) show expansions of coniferous forests during glacial phases rather than steppe vegetation, while interglacials favoured thermophilous forest types. The Lamone marine succession exhibits three forest formations, including mixed-deciduous, Juglandaceae, and mountain coniferous forests, indicating fluctuating humidity levels. The flora experienced significant depletion, particularly with the decline of key taxa (e.g., "Taxodiaceae", *Cathaya*) around 1.2 Ma, marking the transition from subtropical ecosystems to deciduous forests in the Mediterranean (Combourieu-Nebout & Vergnaud Grazzini, 1991; Combourieu-Nebout et al., 2015).

Climate reconstructions reveal a pronounced cooling trend between 1.4 and 1.3 Ma, with reductions in winter temperatures and annual precipitation during both interglacial and glacial phases (Combourieu-Nebout et al., 2015), signalling a progressive increase in aridity, particularly during glacial periods.

### Chbianian (Figs 1-2 and 5-6)

During the Chianian (MIS 19-MIS 6/5), which includes the upper portion of the EMPT (Mudelsee & Schulz, 1997; Clark, 2006; Elderfield et al., 2012), the Mediterranean region continued to experience progressive floristic impoverishment, aligned with climatic gradients, marked by severe glacial episodes (e.g., MIS 16, 12 and 6). The previously dominant 41 kyr G/I cycles (Pisias & Moore, 1981) transitioned to the longer 100 kyr cycles,
extending the duration of glacial stages (Tzedakis, 2005, 2007; Leroy, 2007; Tzedakis et al., 2012) and the pronounced asymmetric pattern of G/I with increasingly “sawtooth” wave form of benthic oxygen isotope stratigraphy (Lourens, 2004; Lisiecki & Raymo, 2005; see also Past Interglacials Working Group of PAGES, 2016). This is the widely studied time interval that also benefits from the high-resolution data and chronology provided by ice cores (e.g., Bazin et al., 2013) and from estimate of sea-level changes (Elderfield et al., 2012). From MIS 11 onwards, a notable rise in interglacial temperatures and CO₂ levels was recorded during the mid-Brunhes Event (MBE), as evidenced by marine, terrestrial, and ice-core data (Masson-Delmotte et al., 2010; Lang & Wolff, 2011; McClymont et al., 2013).

**MARINE RECORDS** - Marine records from different intervals of the Chibanian are well studied in the Mediterranean area based on multiproxy studies, testifying the occurrence of very high amplitude oscillations in climate proxies. The high climate variability influenced the calcareous plankton evolution although no significant turnover occurred in both foraminifera and nannofossil assemblages. They, however, display prominent abundance fluctuations of selected key taxa in response to variations in the environmental conditions at orbital and millennial to centennial scale that were a result of high-low icesheet volume and its instability, strength of thermohaline circulation and of Mediterranean-Atlantic water exchange, and brief influx of polar-subpolar taxa into the Mediterranean. The main examples of such events are from on land marine record of MJS (Marino et al., 2020, see detail below), deep-sea cores in MIS 20-19 transition (TIX, Quivelli et al., 2020, 2021; Toti et al., 2020) and the mid-Brunhes interval (Kandiano et al., 2012; Cortina et al., 2013; Girone et al., 2013b; Maiorano et al., 2013b, 2016b; Milker et al., 2013; Regattieri et al., 2015, 2019; Capotondi et al., 2016; Marino et al., 2018; Trotta et al., 2019; Azibeiro et al., 2021). The latter includes the exceptional cold glacial MIS 12 with sea level about 125 m below present (Bintanja et al., 2005; Elderfield et al., 2012) and the warm long-lasting MIS 11, considered an analogue of the Holocene (de Abreu et al., 2005), with sea level 20 m above the mean present level (e.g., Kindler & Hearty, 2000). The last occurrence of the extinct nannofossil *Pseudoemiliania lacunosa* (Kamptner, 1963) in MIS 12 (e.g., Raffi et al., 2006; Maiorano et al., 2016b) could represent a response to the severe cooling at this time, testified by the incursion of polar calcareous plankton taxa (*N. pachyderma* left coiled and *Coccolithus pelagicus* ssp. *pelagicus* (Wallich, 1877), together with cold low saline melt waters from the Atlantic Ocean into the Mediterranean through the Gibraltar Strait during the southward migration of the Polar Front. The occurrence of these taxa in the Mediterranean Sea allowed to recognise short-term colder events referred as Heinrich-type events (Girone et al., 2013b; Marino et al., 2018; Azibeiro et al., 2021) since their chronology correlates with that of ice rafted detritus (Stein et al., 2009; Alonso-Garcia et al., 2011; Naafs et al., 2011) and lower sea surface temperature (Rodrigues et al., 2011, 2017) in the North Atlantic Ocean, and more arid condition in the Mediterranean region (Larrasoña et al., 2003). The evolution towards increased size (> 4 µm) of nannofossil *Gephyrocapsa margereli*

*muellerae* (sensu Baumann & Freitag, 2004) starting from MIS 12 and MIS 10, has been also recognised as a signal of colder glacials (Girone et al., 2013b; Maiorano et al., 2015; Marino et al., 2018). Whereas the influx of extinct warm nannofossil *Helicosphaera inversa* (Gartner, 1977) distinctly marks MIS 11 in the Mediterranean Sea (Maiorano et al., 2013a), in line with the increase of sea surface temperature and warm water taxa within planktonic foraminifera assemblages (Girone et al., 2013b; Maiorano et al., 2013b, 2016b; Capotondi et al., 2016) during enhanced northward transport of tropical waters in the Atlantic Ocean, as recorded west of Portugal (Voelker et al., 2010) and its influx into the Mediterranean.

**TERRESTRIAL RECORDS** - Pollen records from southern sites during the early Chibanian indicate a dominance of *Artemisia* and *Ephedra* steppes, with alternating temperate to warm-temperate deciduous forests. Significant expansions of steppe taxa were observed at MJS during glacials and stadials within MIS 19 and the base of MIS 16 (Bertini et al., 2015; Toti & Bertini, 2018; Nomade et al., 2019; Toti et al., 2020). Conversely, the Valle di Manche site features alpine-type forests during MIS 18, characterised by *Picea*, *Abies*, and *Fagus* (Capraro et al., 2005, 2015). At Piânico-Sèllere in northern Italy, evidence of a prolonged interglacial phase within MIS 19 is indicated by the expansion of broad-leaved deciduous forests, followed by a contraction of thermophilous species and the rise of coniferous forests (Ravazzi, 2003; Pini et al., 2014). The floristic assemblages reflect climatic fluctuations, with taxa such as *Hippophaë rhamnoides* (Linnaeus, 1753) expanding during colder steppe phases and thermophilous taxa like *Cistus* declining (Suc et al., 1995). Chronological distribution studies have traced the last occurrences of several key taxa, including “Taxodiaceae,” *Carya*, *Pterocarya*, and *Tsuga* (e.g., Bertini, 2010; Combourieu-Nebout et al., 2015; Magri et al., 2017). In northern Italy, *Tsuga’s* presence decreased significantly during the Early-Middle Pleistocene (Piânico-Sèllere: Pini et al., 2014; Moscariello et al., 2000), while it remained in southern sites until MIS 18 (Valle di Manche: Capraro et al., 2005; Montalbano Jonico: Bertini et al., 2015; Toti & Bertini, 2018), though always in low percentages. *Tsuga’s* decline has been linked to increasing global aridity and temperature reductions associated with the shift from 41 to 100 kyr Milankovitch cycles (Bertini, 2000), as well as to potential catastrophic fires during drought periods (Ravazzi et al., 2005). Overall, pollen data indicate that drought intensified during longer glacial periods, with shorter interglacials beginning at cooler temperatures (Combourieu-Nebout et al., 2015).

## LAND-SEA CORRELATIONS IN THE MEDITERRANEAN REGION: CASE STUDIES FROM THE GELASIAN, CALABRIAN AND CHIBANIAN STAGES

### *The Gelasian: the effects of Northern Hemisphere Glaciations (southern Italy) as recorded in the Semaforo section (Figs 1-4 and 7)*

The Semaforo section together with Vrica and Santa Lucia is part of the Crotone series (southern Italy), one of the most thoroughly studied Early Pleistocene succession.
CROTONE
SERIES

Lithology

a

b

c

3

SANTA LUCIA

<figure><img src="image_3.png" /></figure>

VRICA B

2

1

SEMAFORO

<figure><img src="image_4.png" /><figcaption>Fig. 4 - (color online) The Crotone series. a) Main sedimentary, chronological, and biostratigraphic constraints. b) Pollen curves of the main representatives of the vegetation. c) Three windows showing significant marine and terrestrial changes pertinent to palaeoenvironmental reconstructions: the Gelasian (Semaforo) (c1), the Gelasian/Calabrian transition (Vrica B) (c2), the Calabrian/Santa Lucia (c3). Lithology: 1: clays; 2: sapropel; 3; ash. Magnetic polarity: 1: normal; 2: reverse. Wwt indicates warm water foraminifera.</figcaption></figure>
<figure><img src="image_5.png" /><figcaption>Fig. 5 - (color online) a) Panoramic view of the badlands outcropping west of Montalbano Jonico town, showing the upper part of the studied succession and the V5-V9 volcaniclastic layers. b) The Ideale section with the main stratigraphic and palaeoenvironmental features.</figcaption></figure>

Its exceptional location and depositional environment (Fig. 3) provided unparalleled opportunities for advancements in biostratigraphy, magnetostratigraphy, cyclostratigraphy, and precise chronological reconstructions (Suc et al., 2010; Cita et al., 2012, and references therein). The sedimentary succession mainly consists of grey claystones, including four main ash beds and 31 brownish sapropel layers, moderately to strongly laminated (Hilgen, 1991; Zijderveld et al., 1991; Combourieu-Nebout, 1995). The biostratigraphy of foraminifera and calcareous nannofossils within the Crotone series exhibits notable heterogeneity and some inconsistencies

remain unresolved. Different approaches (e.g., Colalongo in Selli et al., 1977; Colalongo et al., 1981; D’Onofrio, 1981; Spaak, 1983; Hilgen, 1990; and unpublished data from J. Cravatte and F. Quillevère, as cited in Joannin et al., 2007) led to some taxonomic inconsistencies and discrepancies in species biostratigraphic ranges, particularly for taxa like *Neogloboquadrina atlantica*. In the case of calcareous nannofossils, early low-resolution studies (e.g., Cati & Borsetti, 1981; Nakagawa et al., 1997) were significantly expanded by Rio et al. (1997), who provided a detailed record of the Vrica section, later complemented by Lourens et al. (1998) using data
from the Santa Lucia section. However, no nannofossil
analyses have been conducted on the Semaforo section.
Other microfossil groups, such as ostracods and diatoms,
along with macrofossils like molluscs and fishes,
offered limited biostratigraphic data (for more detailed
discussions, refer to Pasini & Colalongo, 1997). The
three main sections are stratigraphically more or less
overlapping, i.e., the Semaforo section is entirely within
the Gelasian Stage, the Vrica (A, B, C) section partly
belongs to the Gelasian, partly to the Calabrian Stage, and
the Santa Lucia section belongs to the Calabrian Stage.
The Crotone series covers the time-interval from 2.47 to
1.21 Ma and displays 30 complete G/I cycles, from MIS
97 to MIS 37. Insolation cycles (i-c) are recorded from
i-c236 to i-c116 by combining lithology (sapropels) and
palynology (amorphous organic matter and abundance
in pollen grains of riparian trees, two indices of anoxic
condition development and runoff intensity, respectively).
The understanding of Early Pleistocene G/I cycles pollen
records was clarified as the response of vegetation to the
strong interaction between precession and obliquity as
analysed along several successive climatic cycles.

The Semaforo section, covering the lower part of the Crotone series, from 2.45 to 2.2 Ma, records the first climate cycles (G/I) driven by the obliquity periodicity (about 41 kyr) (Figs 3 and 4b, c1), linked to the onset of northern polar ice sheet. During interglacials, continental areas were dominated, as indicated by pollen data (Combourieu-Nebout, 1993; Fig. 4b, c1), by extensive *Sequoia* forest, and exhibit the recurrent development of subtropical forests associated to warm and yearly humid climate. In contrast, glacials were characterised by the expansion of steppe vegetation and the growth of coniferous forests at higher altitude as evidenced by increases of *Artemisia* and *Cedrus* pollen grains, respectively. These patterns indicate cyclic dry and/or cool conditions. In the marine environment, oxygen isotope curves illustrate the regular alternation between warm and cold conditions, with the earliest G/I alternations detailed in Fig. 4c1. These cycles are characterised by the last occurrence in MIS 96 of cold water *Neogloboquadrina atlantica*, following its short-term pulses in the Mediterranean Sea during colder glacials MIS 100 and MIS 98 as recorded in the Singa section (Zachariasse et al., 1990; Lourens et al., 1992) and Capo Rossello (Caruso, 2004) in agreement with colder sea surface temperature (Herbert et al., 2015, 2018), and by the extinction of *Globorotalia bononiensis* (Dondi, 1962) at the base, as well as the first occurrence of *Globorotalia inflata* (d'Orbigny, 1839a) (e.g., Spaak, 1983). At Capo Rossello, the cyclostratigraphic and micropalaeontological study of Caruso (2004) reports a rich planktonic foraminifera assemblage characterised by fluctuating abundances of warm, temperate and cold taxa in relation to the Gelasian climate cycles, e.g., *Globigerinoides ruber* (d'Orbigny, 1839b), *Trilobatus sacculifer* (Brady, 1877), *Globorotalia crassaformis* (Galloway & Wissler, 1927), *Truncorotalia truncatulinoides* (d'Orbigny, 1839a), *Globigerina bulloides* (d'Orbigny, 1826), and *Turborotalita quinqueloba* (Natland, 1938). Several peaks of *N. atlantica* are observed and correlate with *Artemisia* steppe phases. Oxygen isotope oscillations show consistent intensity, except for MIS 94, which appears less cold, and corresponds to a smaller retreat of forests on the

continent (Combourieu-Nebout, 1993). The selected time
slice of the base of the Semaforo (Fig. 4c1) highlights
the G/I cycle corresponding to MIS 96-95 oscillations.
This interval captures the peak development of *Sequoia*
forest in the Semaforo section and the initial retreat of the
subtropical taxa during the Early Pleistocene. In pollen
record, a decrease in *Sequoia* forest is accompanied by an
increase in *Artemisia* pollen, indicating steppe expansion
during the glacial period (Fig. 4c). Simultaneously, the
occurrence of *Neogloboquadrina atlantica* is a signal
of cooling of sea surface water in the Ionian Sea during
this glacial episode. The warming in the sea water (wwt)
as expressed by the lower value in δ¹⁸O<sub>G. bulloides</sub> and the
*Sequoia* forest pollen record show parallel patterns
illustrating simultaneous temperature variations in the sea
surface and terrestrial environments (Fig. 4c). However,
the wwt curve from planktonic foraminifer δ¹⁸O appears
to rise earlier than the vegetation shift marked by the
expansion of the *Sequoia* forest. The end of the MIS 96
is earlier in the Mediterranean waters suggesting that
response of marine environment to climate changes was
more rapid than continental ones. This is probably due to
the dynamics of the vegetation that need time to get started
after the glacial. During the MIS 95 climatic optimum,
a slight oscillation is observed between *Sequoia* and
*Cathaya* representation in the forest. This might reflect
a minor cooling, as *Cathaya* is known to persist under
cooler conditions.

Calabrian: Intensification of glaciations and
disappearance of subtropical ecosystems in the southern
Mediterranean as recorded in the Vrica and Santa Lucia
sections (southern Italy) (Figs 1-4 and 7)

The Crotone series provides significant and unique stratigraphic and palaeoenvironmental insights also for the Calabrian, as demonstrated by records from the Vrica and Santa Lucia sections (Figs 3 and 4). They document the successive steps of changes in flora and vegetation. The decline and/or disappearance of all subtropical taxa such as *Sequoia*, *Engelhardia*, *Distylium*, and other Hamamelidaceae represent the most prominent event straddling the Gelasian/Calabrian transition. This shift is associated to a notable reduction in flora and ecosystem diversity in southern Italy. The cooling shift is also recorded through the Gelasian-Calabrian transition in marine realm marked by several events that could represent a turnover in calcareous plankton: the last occurrence of *Discoaster brouweri*, the first occurrences of *Globigerina cariacoensis* (Rögl & Bolli, 1973), *Globigerina calabra* (Colalongo & Sartoni, 1977) and of *Globigerinoides tenellus* (Parker, 1958), the extinction of *Globigerinoides obliquus extremus* (Bolli & Bermudez, 1965), the re-occurrence of abundant *Neogloboquadrina pachyderma* left coiled (First Common Occurrence) and the appearance of nannofossil medium-sized *Gephyrocapsa* (4-5.5 µm), followed by the disappearance of *Calidiscus macintyrei* (Bukry & Bramlette, 1969) and the appearance of large *Gephyrocapsa* (> 5.5 µm) (see Cita et al., 2008, 2012; Lirer et al., 2019 and references therein). The increasing size in the placoliths of *Gephyrocapsa* may have been a response to a lowering trend of sea water temperature, in agreement with Kulhanek (2009), Girone et al. (2013b), Marino et al. (2014).
This is shown primarily by the consistent decline in *Sequoia* forest during interglacials, culminating in its near-complete disappearance at ~1.3 Ma in the Santa Lucia section. The concomitant expansion of *Quercus* forest between 2.1 and 1.95 Ma, followed by *Cathaya* forest between ~1.9 and ~1.7 Ma suggests a gradual cooling of interglacial maxima. Strong glacials, as depicted in the oxygen isotope record, occurred just before the expansion of *Cathaya* forest and may have been a contributing factor to the significant retreat of *Sequoia* in southern Italian forests. This decline likely reflects the combined effects of climatic cooling and competition among plant taxa. After 1.7 Ma, the forest underwent a significant restructuring, marked by the rapid retreat of *Cathaya* forest in southern Italy. This is followed by the establishment of a *Quercus* forest with *Carya* at mid-altitude and then of a coniferous forest with *Tsuga* at higher altitude, indicating further cooling of interglacial conditions. Simultaneously, marine evidence in the Vrica section shows the first occurrence of *Hyalinea balthica*, a foraminifer associated with cold waters (Selli, 1967). The oxygen isotopes record strong glacial conditions, especially in sea surface waters, as documented in the Santa Lucia section. Notably, a slight oscillation is observed during the MIS 41 (details in Fig. 4), between *Quercus* and *Tsuga*, likely reflecting a brief temperature drop, with *Tsuga* persisting under cooler conditions. In the Santa Lucia section, the MIS 41 optimum is clearly represented by the abundance of *Quercus* and *Tsuga* and correlated with warming in the sea surface. This warming appears to have commenced earlier in the last G/I cycle recorded in the Vrica section. In parallel, glacials became increasingly severe, marked by the expansion of *Artemisia* steppe, especially in the Santa Lucia section. This reflects enhanced dryness during glacials. In fact, *Artemisia* steppe reaches its maximum extent within the Crotone series at the top of the Santa Lucia section, just after 1.3 Ma, corresponding to an intense glacial period as indicated by oxygen isotope records.

*Chibanian: The climate variability as recorded in the Montalbano section towards the transition to 100,000 year glacial cycles (Figs 1-2 and 5-7)*

The Chibanian is well represented in the Mediterranean region by pelagic-hemipelagic sediments, which predominantly outcrop at several southern Italy sites. Among these, the Ideale section (IS), as a part of the MJS (Ciaranfi et al., 2010; Maiorano et al., 2010) (Fig. 5), holds particular significance. Recently ratified as a Standard Auxiliary Boundary Stratotype (SABS) for the GSSP of the Middle Pleistocene Subseries/Subepoch (Marino et al., 2024) (Fig. 6), it provides a key Mediterranean

reference section for this interval. It is important to highlight that the primary GSSP for the Chibanian Stage was established at the Chiba section in Japan (Suganuma et al., 2021; https://stratigraphy.org/gssps/chibanian). The IS, however, complements this designation, offering a unique perspective from the Mediterranean area, where its stratigraphic features contribute to understanding regional expressions of global climatic and environmental changes during the Middle Pleistocene.

The MJS, encompassing MIS 37-16, is ca. 450 m thick and consists of a coarsening-upwards sedimentary succession ranging from silty clays to silty sands, belonging to the “Argille subappenine” informal unit (Azzaroli et al., 1968). It deposited in the southern Apennines foredeep in a lower circalittoral-upper bathyal setting, which registered even slight climatically induced modifications of local and global significance such as changes in sea level, precipitation, inorganic/organic and fresh water input from land, and tectonic and volcanic events in the Italian Peninsula (D’Alessandro et al., 2003; Stefanelli, 2003, 2004; Maiorano et al., 2008, 2010, 2016a; Ciaranfi et al., 2010; Girone et al., 2013a; Petrosino et al., 2015; Gallicchio et al., 2023). Nine tephra (V1-V9) have been identified, some of which (V3, V4, and V5) were ³⁹Ar/⁴⁰Ar dated (Fig. 5). The MJS is a prime example of multidisciplinary palaeoenvironmental and chronostratigraphic research, which took advantage from terrestrial (pollen grains) (Joannin et al., 2008; Bertini et al., 2015; Marino et al., 2015; Toti, 2015; Maiorano et al., 2016a) and marine (dinoflagellate cysts, calcareous plankton and benthos, ostracods, molluscs, otoliths) fossil proxies (D’Alessandro et al., 2003; Stefanelli, 2003, 2004; Maiorano et al., 2004, 2008, 2010, 2016a, 2021; Ciaranfi & D’Alessandro, 2005; Girone, 2005; Stefanelli et al., 2005; Girone et al., 2013a; Aiello et al., 2015), and from geochemical and biomarker analyses (stable oxygen and carbon isotopes, ¹⁰Be/¹⁹Be, alkenones; Ciaranfi et al., 2010; Simon et al., 2017; Nomade et al., 2019; Marino et al., 2020; Maiorano et al., 2021). The multiproxy approach has provided accurate calcareous plankton (foraminifera and nannofossils) biostratigraphy, recognition of G/I cycles and stadial-interstadial phases, palaeodepth variation, and evidence of sapropel insolation cycles (i-c) 112-74 (Fig. 6).

The Pollen Temperature Index (PTI), defined as the ratio of thermophilous arboreal to steppe taxa, serves as a reliable indicator of regional temperature changes and, when integrated with oxygen isotope curves, enables the identification of glacial and interglacial intervals, as well as submillennial-scale fluctuations, as already observed in the Crotone series (Fig. 4a). Reconstructed for the entire MJS (Bertini et al., 2015; Toti, 2015; Vannacci, 2016),

Fig. 6 - (color online) On the left, the Montalbano Jonico composite section, with the main sedimentary, chronological, biostratigraphic and palaeoenvironmental constraints. LO: lowest occurrence; LCO: lowest common occurrence; HCO: highest common occurrence. On the right, a detail of the Ideale section showing the most important marine and terrestrial record for palaeoenvironmental reconstruction. Data are from Ciaranfi et al. (2010); Maiorano et al. (2010, 2016a); Bertini et al. (2015); Marino et al. (2015, 2020); Petrosino et al. (2015); Simon et al. (2017); Nomade et al. (2019). PTI (Pollen Temperature Index) curve is according to Bertini et al. (2015); Toti et al. (2015); Vannacci et al. (2016). In the Ideale section: end td2 is the end of the temporary disappearance of *G. omega*; Total N indicates total nannofossil absolute abundance; wwt indicates warm water taxa; MF is the maximum flooding; MD is the maximum depth; I, II and III are the three warmer phases in MIS 19c; star symbol is the abundance peak of *P. zoharyi*; Med-H is the terminal stadial-like event; Med-BA is the Bølling-Allerød-like event; Med-YD is the Younger Dryas-like episode.
Montalbano Jonico
composite section

Biozone

Pseudoemiliania lacunosa

small Gephyrocapsa

<figure><img src="image_6.png" /></figure>

Ideale section

<figure><img src="image_7.png" /></figure>
<figure><img src="image_8.png" /></figure>

<figure><img src="image_9.png" /></figure>

Mediterranean
taxa

<figure><img src="image_10.png" /></figure>

Ericaceae

<figure><img src="image_11.png" /></figure>

Steppic and
dry herbs

<figure><img src="image_12.png" /></figure>

Other herbs

<figure><img src="image_13.png" /></figure>

Gephyrocapsa
omega

<figure><img src="image_14.png" /></figure>

Umbilicosphaera
sibogae

<figure><img src="image_15.png" /></figure>

Globorotalia
crassaformis

<figure><img src="image_16.png" /></figure>

Truncorotalia
truncatulinoides

<figure><img src="image_17.png" /></figure>

Globorotalia
inflata

<figure><img src="image_18.png" /></figure>

Globigerinoides
ruber

<figure><img src="image_19.png" /></figure>

Trilobatus
sacculifer

<figure><img src="image_20.png" /></figure>

Discoaster
brouweri

<figure><img src="image_21.png" /></figure>

Calcidiscus
macintyrei

<figure><img src="image_22.png" /></figure>

Gephyrocapsa
oceanica

<figure><img src="image_23.png" /></figure>

large
Gephyrocapsa

<figure><img src="image_24.png" /></figure>

Helicosphaera
sellii
the PTI reveals distinctive oscillations between warmer and colder phases that closely align with variations in palaeodepth (see fig. 2 in Marino et al., 2024) and G/I patterns (Fig. 6). Notably, the amplitude variations in the PTI curve, in conjunction with the benthic δ¹⁸O, are particularly striking. These covariant changes, observed between the lower and upper parts of the MJS, reflect the progressive establishment of the 100 kyr periodicity of G/I cycles, beginning from MIS 22 onward.

The IS (Figs 5-6), ca 80 m thick, covers the time interval from MIS 20 to the onset of MIS 18. The high temporal resolution study of this section (up to 200 years; Nomade et al., 2019) analysed the same samples based on pollen and calcareous plankton assemblages, carbon and oxygen isotopes together with alkenones, the latter providing the alkenone-derived sea surface temperature (alkenone-SST) and the total C₃₇ as a proxy of marine palaeoproductivity (Marino et al., 2020; Maiorano et al., 2021). G/I andstadial/interstadial phases have been recognised in agreement with δ¹⁸O, δ¹³C, Mesothermic Arboreal Pollen Taxa (MAPT) and calcareous plankton dataset. The signature of orbital to millennial climate variability is clearly imprinted in the IS as recorded by multiproxy investigations and by the alternating dark grey and light grey sediment color exposed in the field, which correspond to the warm-wet and cold-dry climate phases, respectively (Fig. 5). In late MIS 20, between 794 and 788.5 ka, a terminal stadial-like event (Med-H<sub>TIX</sub>, Med-H in Fig. 6) occurred as a result of combined atmospheric dynamics and oceanic/ hydrological processes (Fig. 6) (Marino et al., 2020). The Med-H<sub>TIX</sub> is characterised by: 1) an alkenone-SST 4-5 °C cooler than in the older interval, 2) higher abundances of arctic-subarctic *Coccolithus pelagicus* ssp. *pelagicus* and *Neogloboquadrina pachyderma* (Maiorano et al., 2016a) and decrease of warm-oligotrophic calcareous plankton taxa (Fig. 6), 3) a large expansion of open landscapes including prevalent (cold) dry steppe on land (Bertini et al., 2015) and a slight decrease of MAPT (Fig. 6), and 4) a lightening of 1‰ in the δ¹⁸O<sub>M.barleeanum</sub> (Nomade et al., 2019) reflecting the influx of lighter fresh water into the Ionian Sea (meltwaters coming from mountain glaciers of the Alpine and Apennines chains or from North Atlantic via the Gibraltar Strait) (Fig. 6). The Med-H<sub>TIX</sub> at the IS is a signal of both a local and global climate episode since it correlates with a peak of C₃₇.₄ (low salinity and colder meltwaters) at the western Mediterranean ODP Site 975 (Quivelli et al., 2021) and at the Iberian Margin Integrated Ocean Drilling Program (IODP) Site U1385 (Rodrigues et al., 2017) as well as with the deposition of ice-rafted debris (IRD) in the northern Atlantic (Wright & Flower, 2002). During MIS 20-19 deglaciation (Termination IX), millennial scale episodes before the onset of full interglacial MIS 19c have been identified and they recall the high frequency climate oscillation of Termination I (see Maiorano et al., 2016a; Marino et al., 2020). In this respect, a warm episode has been referred to a Bølling-Allerød-like event (Med-BA<sub>TIX</sub>, Med-BA in Fig. 6),

characterised by a climate amelioration (alkenone-SST increase of 4 °C, slight increase of warm water calcareous plankton taxa and MAPT) also documented by the increase of dinoflagellate cysts *Spiniferites mirabilis/hyperacanthus* (Rossignol, 1964; Sarjeant, 1970) and distinct patterns of *Orbulina universa* (d'Orbigny, 1839b) and *Globorotalia inflata*. Upwards, a Younger Dryas-like episode (Med-YD<sub>TIX</sub>, Med-YD in Fig. 6) has been suggested based on a 2.5 °C decrease of alkenone-SST, the short-term peaks of *Coccolithus pelagicus* ssp. *pelagicus*, *Neogloboquadrina pachyderma* and *Neogloboquadrina incompta* (Cifelli, 1961), and slight decrease of warm water calcareous plankton taxa (wwt in Fig. 6) and MAPT (Fig. 6). The combined analyses were able to detect the ghost sapropel i-c74 in early MIS 19c, from 785.0 ka to 782.6 ka and its internal variability has been described in detail (Maiorano et al., 2016a; Nomade et al., 2019; Marino et al., 2020). Together with the δ¹³C minimum (water column stratification, higher organic matter at the sea floor), an increase in the benthic foraminiferal infauna/ epifauna ratio (sea bottom stress; Stefanelli, 2003) and a peak of the dinoflagellate cyst *Polysphaeridium zoharyi* (Rossignol, 1962) (less saline water; Bertini et al., 2015; Maiorano et al., 2016a) (star in Fig. 6) showed an increase during i-74. It is worth noting that the interruption of the stable and stratified surface water conditions in the middle of sapropel evidenced by calcareous plankton as a signal of a short-term increase in nutrients and mixing in sea water (Marino et al., 2020) is accompanied by a brief cooler and dryer episode on land indicated by a decrease in the MAPT curve (Fig. 6), suggesting a synchronous response of terrestrial and marine realms to even millennial climate changes. The warmer climate conditions and the climate optimum during MIS 19c lasted for ~11.5 ± 3.4 kyr (Nomade et al., 2019) similarly to the full interglacial recorded in the lacustrine Sulmona section (i.e., 11 ky; Regattieri et al., 2019) or inferred in the North Atlantic records (i.e., 12.5-10.5 ky; Tzedakis et al., 2012; Head, 2021). This warm phase was marked by a distinct increase of calcareous plankton warm taxa, such as *Umbilicosphaera sibogae* (Weber van Bosse, 1901), *G. caribbeanica*, *G. omega*, *Oolithotus* spp., *Globigerinoides ruber*, and *Trilobatus sacculifer*. A contemporary response to marine and terrestrial environmental variation is highlighted by the three warmer phases visible during the full interglacial MIS 19c, as recorded by the patterns of total nannofossil absolute abundance (Total N in Fig. 6) and warm water calcareous nannofossil taxa, in concert with the increase of alkenone-SST and of MAPT. Three main increases of the Mediterranean Forest Pollen during Tajo phase (Sánchez-Goñi et al., 2016) were also detected on the Iberian margin at the IODP site U1385 similarly to what occurred in the MIS 19c at the IS highlighting a shared response of vegetation to atmospheric processes that, in concert to repeated tropical subtropical water influxes through the Gibraltar Strait towards the central Mediterranean Sea, also influenced the marine proxies (Marino et al., 2020). It is remarkable to observe how the

Fig. 7 - (color online) Major environmental shifts in terrestrial and marine ecosystems of southern Italy during selected time windows. Key changes in both terrestrial vegetation and marine conditions illustrate the dynamic interplay between land and sea from MIS 95 to MIS 19c. On the left is the Standard Biochronostratigraphy, see Fig. 2 for the legend and references. Micrograph images of selected taxa are not to scale.
upwardstadial-interstadial oscillations in MIS 19b-MIS
19a interval toward the MIS 18 onset are synchronously
recorded by both MAPT and calcareous plankton data
set, which together with the δ¹⁸O, δ¹³C and alkenone-
SST patterns indicate short-term changeable conditions
in marine and terrestrial environments. In this respect:
1) warmer and wetter on land climate corresponds to
enhanced chemical on land weathering and continental
fresh water input (Bertini et al., 2015; Maiorano et al.,
2016a; Nomade et al., 2019), and stratified sea surface
waters during higher sea level within interstadials (MIS
19a-1 and 19a-3 in Fig. 6) (D'Alessandro et al., 2003;
Stefanelli, 2003; Aiello et al., 2015; Bertini et al., 2015;
Marino et al., 2015; Maiorano et al., 2021), while 2) colder
and drier terrestrial climate occurred when sea surface
waters experienced lower temperature and mixing during
lower sea level within stadials.

CONCLUDING REMARKS

This synthesis integrated Italian pollen records with marine datasets (Figs 1 and 2), providing a comprehensive view of both terrestrial and marine environments during the latest Pliocene and Pleistocene (Figs 3-7) and improving the timing of events and changes in a shared chronology. Pollen data from the Mediterranean region reveal significant shifts in vegetation, which align with glacial-interglacial cycles, stadial/interstadial phases, and rapid, short-lived events. As summarised in Bertini et al. (2010) and Bertini & Combourieu-Nebout (2023), the glacial-interglacial cycles are generally characterised by the predominance of steppe communities during glacials, marked by the expansion of *Artemisia* and *Ephedra*, indicating progressive establishment of cooler and drier conditions. In contrast, interglacial phases support broadleaf deciduous forests, reflecting favourable climatic conditions for diverse flora. The alternation between cold and warm phases during the Late Pliocene and Early Pleistocene influenced the evolution of calcareous plankton in marine environment as evidenced by appearance and extinction of climate-sensitive taxa (e.g., last occurrence of *Discoaster brouweri*) or their abundance increase and temporary disappearance (e.g., *Neogloboquadrina pachyderma* left coiling), and short-term influxes (e.g., *N. atlantica*) of polar key species in the Mediterranean Sea. The transition from 41 kyr to 100 kyr glacial cycles profoundly impacted both terrestrial and marine ecosystems determining the gradual disappearance of subtropical taxa and ecosystems (e.g., Bertini, 2010; Magri et al., 2017; Bertini & Combourieu-Nebout, 2023 and references therein). This is particularly well-documented in the sites selected in this paper (Figs 4, 6 and 7), starting with the reduction and eventual disappearance of *Sequoia* in the upper portion of Semaforo, followed by the reduction of *Cathaya* and subsequently *Tsuga*, especially evident in the sites of Vrica-Santa Lucia and MJS (Fig. 7). Previous palynological data also contributed to describing the presence of ecological and climatic gradients across the region, at least since the latest Pliocene. The calcareous plankton experienced a turnover marked by extinction and appearance of species in the Vrica series and recorded significant orbital-suborbital

changes in the MJS, where variation in the abundance of
warm and cold taxa with different ecological preference
for nutrients and photic column water structure have been
detected in line with climate-induced change in terrestrial
environments. From approximately MIS 43 to MIS 16,
the quite well-calibrated Italian pollen records can be
compared with key reference sites across Europe, such
as Lake Ohrid (Donders et al., 2021, 2024) and Tenaghi
Philippon (e.g., Pross et al., 2015), as well as those
from the Iberian margin (Sánchez-Goñi et al., 2023 and
references therein), which offer long, continuous archives
that enhance our understanding of floristic and climatic
dynamics. Integrating these records has enabled us to
situate Italian successions within a broader European
framework, providing critical insights into vegetation
shifts and climate oscillations at regional and global
scales. Specifically, the Ohrid data document temperate
and montane forest dynamics, as well as the persistence
and subsequent retreat of subtropical taxa, which align
with the patterns observed in the Italian sites (Fig. 7).

By harmonising terrestrial and marine records, we
captured the intricate feedback between oceanic and
continental systems. The three selected case studies
from Semaforo, Vrica-Santa Lucia, and Montalbano
Jonico illustrate how shifts in climate drivers influenced
both vegetation and calcareous plankton. These
findings emphasise the Mediterranean’s dual role as
both a recorder of global climate variability and a
driver of regional ecosystem changes. This integrated
approach has allowed us to refine chronologies, cross-
validate climatic signals, and develop a more nuanced
understanding of ecosystem responses to orbital- and
sub-orbital-scale climate forcing. The marine record,
including the deep-sea cores recovered by ODP
expeditions (Figs 1 and 2), offers a parallel narrative,
revealing changes in sea surface temperature, salinity,
and ocean circulation patterns that correlate with changes
in terrestrial environment. These marine sites underscore
the interconnectedness of land-sea systems, allowing for
a comprehensive understanding of palaeoenvironmental
conditions. The synchronicity observed in marine
proxies, such as planktonic foraminifera and calcareous
nannofossils, reinforces the idea that climatic changes
were the expression of complex atmospheric and oceanic
dynamics.

Unravelling land-sea correlations during the Pliocene-Pleistocene in the Mediterranean, this contribution adopts a multidisciplinary approach that combines palynological data with marine microfossils, providing crucial insights into climate variability and ecosystem dynamics. These studies are also vital for tracing faunal histories, building on the pioneering work of Augusto Azzaroli, Danilo Torre, and Giovanni Ficcarelli, whose legacy continues to inspire interdisciplinary research. We are honoured to dedicate this work to their memory.

ACKNOWLEDGEMENTS

We would like to thank the editors for their valuable assistance throughout the review process. Our sincere appreciation also goes to Prof. Antonio Caruso and an anonymous reviewer for their insightful comments and constructive suggestions, which greatly contributed to the improvement of this article.
REFERENCES

Addante M., Bertini A., Girone A., Maiorano P., Marino M., Niccolini G., Scopelliti G., Tabbabi H. & Caruso A. (2024a). Climate variability and biotratigraphic constraints at the Neogene-Quaternary boundary at the Monte San Nicola type-section, Sicily. *37th International Geological Congress, Busan, South Korea, Abstract book* : 338-339.

Addante M., Maiorano P., Scopelliti G., Girone A., Marino M., Trotta S. & Caruso A. (2024b). Climate-induced surface water variability at Monte San Nicola type-section (Sicily, southern Italy): New data across the Gelasian GSSP. *Palaeogeography, Palaeoclimatology, Palaeoecology*, 634: 111907.

Addante, M., Herbert, T.D., Girone, A., Caruso, A., Marino, M., Scopelliti, G., Fasone, S. & Maiorano, P. (2025). Monsoon variability and high latitude climate signals in the Central Mediterranean at the Pliocene - Pleistocene transition: The Gelasian Stratotype section (Monte San Nicola, Sicily). *Global and Planetary Change*, 249 (2025) 104788. https://doi.org/10.1016/j.gloplacha.2025.104788

Aiello G., Barra D. & Parisi R. (2015). Lower-Middle Pleistocene ostracod assemblages from the Montalbano Jonico section (Basilicata, southern Italy). *Quaternary International*, 383: 47-73.

Alonso-Garcia M., Sierro F.J. & Flores J.A. (2011). Arctic front shifts in the subpolar North Atlantic during the Mid-Pleistocene (800-400 ka) and their implications for ocean circulation. *Palaeogeography, Palaeoclimatology, Palaeoecology*, 311: 268-280.

Ausín B., Flores J.A., Sierro F.J., Bárcena M.A., Hernández-Almeida I., Frances G., Gutierrez-Arnillas E., Martrat B., Grimalt J.O. & Cacho I. (2015). Coccolithophore productivity and surface water dynamics in the Alboran Sea during the last 25 kyr. *Palaeogeography, Palaeoclimatology, Palaeoecology*, 418: 126-140.

Azibeiro L.A., Sierro F.J., Capotondi L., Lirer F., Andersen N., González-Lanchas A., Alonso-Garcia M., Flores J.-A., Cortina A. & Grimalt J.O. (2021). Meltwater flux from northern ice-sheets to the Mediterranean during MIS 12. *Quaternary Science Review*, 268: 107108.

Azzaroli A., Perno U. & Radina B. (1968). Note illustrative della Carta Geologica d'Italia alla scala 1:100.000, Foglio 188 Gravina di Puglia. 57 pp. Istituto Poligrafico e Zecca dello Stato, Roma.

Bárcena M.A., Cacho I., Abrantes F., Sierro F.J., Grimalt J.O. & Flores J.A. (2001). Paleoproductivity variations related to climatic conditions in the Alboran Sea (western Mediterranean) during the last glacial-interglacial transition: the diatom record. *Palaeogeography, Palaeoclimatology, Palaeoecology*, 167: 337-357.

Bartoli G., Sarnthein M., Weinelt M., Erlenkeuser H., Garbe-Schönberg D. & Lea D.W. (2005). Final closure of Panama and the onset of northern hemisphere glaciation. *Earth and Planetary Science Letters*, 237: 33-44.

Bartoli G., Hönisch B. & Zeebe R.E. (2011). Atmospheric CO₂ decline during the Pliocene intensification of Northern Hemisphere glaciations. *Paleoceanography*, 26: PA4213.

Baumann K.H. & Freitag T. (2004). Pleistocene fluctuations in the northern Benguela Current system as revealed by coccolith assemblages. *Marine Micropaleontology*, 52: 195-215.

Baumann K.H., Andruleit H., Böckel B., Geisen M. & Kinkel H. (2005). The significance of extant coccolithophores as indicators of ocean water masses, surface water temperature, and palaeoproductivity: a review. *Paläontologische Zeitschrift*, 79: 93-112.

Bazin L., Landais A., Lemieux-Dudon B., Toyé Mahamadou Kele H., Veres D., Parrenin F., Martinerie P., Ritz C., Capron E., Lipenkov V., Loutre M.-F., Raynaud D., Vinther B., Svensson A., Rasmussen S.O., Severi M., Blunier T., Leuenberger M., Fischer H., Masson-Delmotte V., Chappellaz J. & Wolff E. (2013). An optimized multi-proxy, multi-site Antarctic ice and

gas orbital chronology (AICC2012): 120-800 ka. *Climate of the Past*, 9: 1715-1731.

Beaufort L., Bassinot F. & Vincent E. (1999). Primary production response to orbitally induced variations of the Southern Oscillation in the Equatorial Indian Ocean. *In Abrantes F. & Mix A.C. (eds), Reconstructing Ocean History*. Springer, Boston, MA: 245-271.

Becker J., Lourens L.J., Hilgen F.J., Van Der Laan E., Kouwenhoven T.J. & Reichart G.J. (2005). Late Pliocene climate variability on Milankovitch to millennial time scales: A high-resolution study of MIS100 from the Mediterranean. *Palaeogeography, Palaeoclimatology, Palaeoecology*, 228: 338-360.

Berggren W.A. (1972). Cenozoic biostratigraphy and paleobiogeography of the North Atlantic. *Initial Reports of the Deep Sea Drilling Project*, 12: 965-1001.

Berner N., Trauth M.H. & Holschneider M. (2022). Bayesian inference about Plio-Pleistocene climate transitions in Africa. *Quaternary Science Reviews*, 277: 107287.

Bertini A. (1994). Messinian-Zanclean vegetation and climate in north-central Italy. *Historical Biology*, 9: 3-10.

Bertini A. (2000). Pollen record from Colle Curti and Cesi: early and middle Pleistocene mammal sites in the Umbro-Marchean Apennine mountains (central Italy). *Journal of Quaternary Sciences*, 15: 825-840.

Bertini A. (2001). Pliocene climatic cycles and altitudinal forest development from 2.7 Ma in the northern Apennines (Italy): evidence from the pollen record of the Stirone section (~5.1 to ca. 2.2 Ma). *Geobios*, 34: 253-265.

Bertini A. (2010). Pliocene to Pleistocene palynoflora and vegetation in Italy: state of the art. *Quaternary International*, 225: 5-24.

Bertini A. (2013). Climate and vegetation in the Upper Valdarno Basin (central Italy) as a response to Northern Hemisphere insolation forcing and regional tectonics in the late Pliocene-early Pleistocene. *II Quaternario*, 132: 137-148.

Bertini A. & Combourieu-Nebout N. (2023). Piacenzian to Late Pleistocene Flora and Vegetation in Italy: A Moving Sketch. *Alpine and Mediterranean Quaternary*, 36: 91-119.

Bertini A., Ciaranfi N., Marino M. & Palombo M.R. (2010). Proposal for Pliocene and Pleistocene land-sea correlation in the Italian area. *Quaternary International*, 219: 95-108.

Bertini A., Toti F., Marino M. & Ciaranfi N. (2015). Vegetation and climate across the Early-Middle Pleistocene transition at Montalbano Jonico, southern Italy. *Quaternary International*, 383: 74-88.

Bertini A., Niccolini G. & Tabbabi H. (2024). Vegetation dynamics and climate changes during the Onset of Northern Hemisphere Glaciation: Insights from the Monte San Nicola Section (central Mediterranean). *37th International Geological Congress, Busan, South Korea, Abstract book*: 333-334.

Bertolani Marchetti D., Accorsi C.A., Pelosi G. & Raffi S. (1979). Palynology and stratigraphy of the Plio-Pleistocene sequence of the Stirone River (northern Italy). *Pollen et Spores*, 21: 149-167.

Bertoldi R., Rio D. & Thunell R. (1989). Pliocene-Pleistocene vegetational and climatic evolution of the south-central Mediterranean. *Palaeogeography, Palaeoclimatology, Palaeoecology*, 72: 263-275.

Bintanja R., van de Wal R.S.W. & Oerlemans J. (2005). Modelled atmospheric temperatures and global sea levels over the past million years. *Nature*, 437: 125-128.

Bolli H.M. & Bermudez P.J. (1965). Zonation based on planktonic foraminifera of middle Miocene to Pliocene warm-water sediments. *Boletin Informativo Asociacion Venezolana de Geologia, Mineria y Petroleo*, 8: 121-149.

Bolton C.T., Wilson P.A., Bailey I., Friedrich O., Beer C.J., Becker J., Baranwal S. & Schiebel R. (2010). Millennial-scale climate variability in the subpolar North Atlantic Ocean during the late Pliocene. *Paleoceanography*, 25: PA4218.

Brady H.B. (1877). Supplementary note on the foraminifera of the Chalk (?) of the New Britain group. *Geological Magazine*, 4: 534-536.
Bramlette M.N. & Riedel W.R. (1954). Stratigraphic value of discoasters and some other microfossils related to Recent coccolithophores. *Journal of Paleontology*, 28: 385-403.

Bukry D. (1973). Coccolith stratigraphy, eastern equatorial Pacific, Leg 16, Deep Sea Drilling Project. *Initial Reports of the Deep Sea Drilling Project*, 16: 653-711.

Bukry D. & Bramlette M.N. (1969). Some new and stratigraphically useful calcareous nanofossils of the Cenozoic. *Tulane Studies in Geology*, 7: 131-142.

Capotondi L., Girona A., Lirer F., Bergami C., Verducci M., Vallefuoco M., Afferri A., Ferraro L., Pelosi N. & De Lange G.J. (2016). Central Mediterranean Mid-Pleistocene paleoclimatic variability and its connection with global climate. *Palaeogeography, Palaeoclimatology, Palaeoecology*, 442: 72-83.

Capraro L. & Maiorano P. (2023). Italian GSSPs of the Quaternary System. *Alpine and Mediterranean Quaternary*, 36: 1-23.

Capraro L., Asioli A., Backman J., Bertoldi R., Channell J.E.T., Massari F. & Rio D. (2005). Climatic patterns revealed by pollen and oxygen isotope records across the Matuyama Brunhes Boundary in the central Mediterranean (southern Italy). *In* Head M.D. & Gibbard P.L. (eds), *Early-Middle Pleistocene Transition: the Land-Ocean Evidence. Geological Society of London Special Publication*, 247: 159-182.

Capraro L., Macrì P., Scarponi D. & Rio D. (2015). The lower to Middle Pleistocene Valle di Manche section (Calabria, Southern Italy): state of the art and current advances. *Quaternary International*, 383: 36-46.

Capraro L., Bonomo S., Di Stefano A., Ferretti P., Fornaciari E., Galeotti S., Incarbona A., Macri P., Raffi I., Sabatino N., Speranza F., Sprovieri M., Di Stefano E., Sprovieri R. & Rio D. (2022). The Monte San Nicola section (Sicily) revisited: A potential unit-stratotype of the Gelasian Stage. *Quaternary Science Reviews*, 278: 107367.

Caruso A. (2004). Climatic changes during late Pliocene and early Pleistocene at Capo Rossello (Sicily, Italy): response from planktonic foraminifera. *In* Coccioni R., Galeotti S. & Lirer F. (eds), *Proceedings of the First Italian Meeting on Environmental Micropaleontology. Grzybowski Foundation Special Publication*, 9: 17-36.

Castradori D., Rio D., Hilgen F.J. & Lourens L.J. (1998). The global standard stratotype section and point (GSSP) of the Piacenzian Stage (Middle Pliocene). *Episodes*, 21: 88-93.

Cati F. & Borsetti A.M. (1981). Calcareous nannoplankton biostratigraphy of the Vrica Section (Calabria, Southern Italy). *Giornale di Geologia*, 43: 365-384.

Channell J., Di Stefano E. & Sprovieri R. (1992). Calcareous Plankton Biostratigraphy, Magnetostatigraphy and Paleoclimatic History of the Plio-Pleistocene Monte San Nicola Section (Southern Sicily). *Bollettino della Società Paleontologica Italiana*, 31: 351-382.

Ciaranfi N. & D'Alessandro A. (2005). Overview of the Montalbano Jonico area and section: a proposal for a boundary stratotype for the lower-middle Pleistocene, southern Italy. *Foredeep. Quaternary International*, 131: 5-10.

Ciaranfi N., Lirer F., Lirer L., Lourens L.J., Maiorano P., Marino M., Petrosino P., Sprovieri M., Stefanelli S., Brilli M., Girone A., Joannin S., Pelosi N. & Vallefuoco M. (2010). Integrated stratigraphy and astronomical tuning of the Lower-Middle Pleistocene Montalbano Jonico land section (southern Italy). *Quaternary International*, 210: 109-120.

Cifelli R. (1961). *Globigerina incompta*, a new species of pelagic foraminifera from the North Atlantic. *Contributions from the Cushman Foundation for Foraminiferal Research*, 12: 83-86.

Cita M.B. (1975). Planktonic foraminiferal biozonation of the Mediterranean Pliocene deep sea record. A revision. *Rivista Italiana di Paleontologia e Stratigrafia*, 81: 527-544.

Cita M.B., Capraro L., Ciaranfi N., Di Stefano E., Marino M., Rio D., Sprovieri R. & Vai G.B. (2006). Calabrian and Ionian: A proposal for the definition of Mediterranean stages for the Lower and Middle Pleistocene. *Episodes*, 29: 107-114.

Cita M.B., Capraro L., Ciaranfi N., Di Stefano E., Lirer F., Maiorano P., Marino M., Raffi I., Rio D., Sprovieri R., Stefanelli S. & Vai G.B. (2008). The Calabrian Stage redefined. *Episodes*, 31: 408-419.

Cita M.B., Gibbard P.L., Head M.J. & the ICS Sub-commission on Quaternary Stratigraphy (2012). Formal ratification of the GSSP for the base of the Calabrian Stage (second stage of the Pleistocene Series, Quaternary System). *Episodes*, 35: 388-397.

Clark P.U. (2006). The middle Pleistocene transition: characteristics, mechanisms, and implications for long-term changes in atmospheric pCO₂. *Quaternary Science Review*, 25: 23-24.

Colalongo M.L. & Sartoni S. (1977). *Globigerina calabra* nuovo specie presso il limite Plio-Pleistocene della Sezione Vrica (Calabria). *Giornale di Geologia, serie 2*, 42: 207-209.

Colalongo M.L., Pasini G. & Sartoni S. (1981). Remarks on the Neogene-Quaternary boundary and the Vrica section (Calabria, Italy). *Bollettino della Società Paleontologica Italiana*, 20: 99-120.

Colleoni F., Cherchi A., Masina S. & Brierley C.M. (2015). Impact of global SST gradients on the Mediterranean runoff changes across the Plio-Pleistocene transition. *Paleoceanography*, 30: 751-767.

Combourieu-Nebout N. (1993). Vegetation response to upper Pliocene glacial/interglacial cyclicity in the central Mediterranean. *Quaternary Research*, 40: 228-236.

Combourieu-Nebout N. (1995). Réponse de la végétation de l'Italie méridionale au seuil climatique de la fin du Pliocène d'après l'analyse pollinique haute résolution de la section de Semaforo (2,46 à 2.1 Ma). *Comptes Rendus de l'Académie des Sciences Paris*, 321: 659-665.

Combourieu-Nebout N. & Vergnaud-Grazzini C. (1991). Late Pliocene northern hemisphere glaciations: the continental and marine responses in the central Mediterranean. *Quaternary Sciences Review*, 10: 319-334.

Combourieu-Nebout N., Semah F. & Djubiantono T. (1990). La limite Pliocène/Pléistocène: précisions magnétostratigraphiques par l'étude sériée de la coupe type de Vrica (Crotone, Italie). *Comptes Rendus de l'Académie des Sciences Paris*, 311: 851-857.

Combourieu-Nebout N., Foucault A. & Mélières F. (2004). Vegetation markers of palaeoclimate cyclical changes in the Pliocene of Punta Piccola (Sicily, Italy). *Palaeogeography, Palaeoclimatology, Palaeoecology*, 214: 55-66.

Combourieu-Nebout N., Bertini A., Russo-Ermolli E., Peyron O., Klotz S., Montade V., Fauquette S., Allen J.R.M., Fusco F., Goring S., Huntley B., Joannin S., Lebreton V., Magri D., Martinetto E., Orain R. & Sadori L. (2015). Climate changes in the central Mediterranean and Italian vegetation dynamics since the Pliocene. *Review of Palaeobotany and Palynology*, 218: 127-147.

Cortina A., Sierro F.J., Filippelli G., Flores J.A. & Berné S. (2013). Changes in planktic and benthic foraminifer assemblages in the Gulf of Lions, off south France: response to climate and sea level change from MIS 6 to MIS 11. *Geochemistry, Geophysics, Geosystems*, 14: 1258-1276.

D'Alessandro A., La Perna R. & Ciaranfi N. (2003). Response of macrobenthos to changes in paleoenvironment in the Lower-Middle Pleistocene (Lucania Basin, southern Italy). *Il Quaternario*, 16: 167-182.

de Abreu F.F., Abrantes N.J., Shackleton P.C., Tzedakis J.F., McManus D.W. & Hall Oppo M.A. (2005). Ocean climate variability in the eastern North Atlantic during interglacial marine isotope stage 11: a partial analogue to the Holocene? *Paleoceanography*, 20: 3009.

De Boer B., Peters M. & Lourens L.J. (2021). The transient impact of the African monsoon on Plio-Pleistocene Mediterranean sediments. *Climate of the Past*, 17: 331-334.

de la Vega E., Chalk T. B., Wilson P. A., Bysani R.P. & Foster G.L. (2020). Atmospheric CO₂ during the Mid-Piacenzian Warm Period and the M2 glaciation. *Scientific Reports*, 10: 1-8.
Donders T., Panagiotopoulos K., Koutsodendris A., Bertini A., Mercuri A.M., Masi A., Combourieu-Nebout N., Joannin S., Kouli K., Kousis I., Peyron O., Torri P., Florenzano A., Francke A., Wagner B. & Sadori L. (2021). 1.36 million years of Mediterranean forest refugium dynamics in response to glacial-interglacial cycle strength. *Proceedings of the National Academy of Sciences, USA*, 118: e202611118.

Donders T.H., Oliveira D., Sánchez-Goñi M.F. & Sadori L. (2024). A 1.1-million-year vegetation history of the Mediterranean region: Scientific drilling reveals sensitivities of forest ecosystems. *Pages Magazine*, 32: 118-119.

Dondi L. (1962). Nota paleontologico-stratigrafica sul pede-Appennino Padano. *Bollettino della Società Geologica Italiana*, 81: 113-230.

D'Onofrio S. (1981). I Foraminiferi bentonici della Sezione Vrica (Calabria, Italia). *Giornale di Geologia*, 43: 327-364.

d'Orbigny A. (1826). Tableau methodique de la Classe de Cephalopodes. *Annales des Sciences Naturelles, Paris*, 7: 245-314.

d'Orbigny A. (1839a). Foraminifères des îles Canaries. *Histoire naturelle des Iles Canaries*, 2: 120-146.

d'Orbigny A. (1839b). Foraminifères. In de la Sagra R. (ed.), Histoire physique et naturelle de l'Ile de Cuba. 224 pp. A. Bertrand, Paris.

Driever B.W.M. (1988). Calcareous nannofossil biostratigraphy and paleoenvironmental interpretation of the Mediterranean Pliocene. *Utrecht Micropaleontological Bulletins*, 36: 1-245.

Dumitru O.A., Austermann J., Polyak V.J., Fornós J.J., Asmerom Y., Ginés J., Ginés A. & Onac B.P. (2021). Sea-level stands from the Western Mediterranean over the past 6.5 million years. *Scientific Reports*, 11: 261.

Elderfield H., Ferretti P., Greaves M., Crowhurst S., McCave I.N., Hodell D. & Piotrowski A.M. (2012). Evolution of ocean temperature and ice volume through the mid-Pleistocene climate transition. *Science*, 337(6095): 704-709.

Fauquette S. & Bertini A. (2003). Quantification of northern Italy Pliocene climate from pollen data: evidence for a very peculiar climate pattern. *Boreas*, 32: 361-369.

Fletcher W.J. & Sánchez-Goñi M.F. (2008). Orbital-and sub-orbital-scale climate impacts on vegetation of the western Mediterranean basin over the last 48,000 yr. *Quaternary Research*, 70: 451-464.

Fusco F. (2007). Vegetation response to early Pleistocene climatic cycles in the Lamone valley (Northern Apennines, Italy). *Review of Palaeobotany and Palynology*, 145: 1-23.

Gallicchio S., Colacicco R., Capolongo D., Girona A., Maiorano P., Marino M. & Ciaranfi N. (2023). Geological features of the Special Nature Reserve of Montalbano Jonico Badlands (Basilicata, Southern Italy). *Journal of Maps*, 19: 2179435.

Galloway J.J. & Wissler S.G. (1927). Pleistocene foraminifera from the Lomita Quarry, Palos Verdes Hills, California. *Journal of Paleontology*, 1: 35-87.

Gartner S. (1977). Calcareous nannofossil biostratigraphy and revised zonation of the Pleistocene. *Marine Micropaleontology*, 2: 1-25.

Girone A. (2005). Response of otolith assemblages to sea-level fluctuations at the Lower Pleistocene Montalbano Jonico section (southern Italy). *Bollettino della Società Paleontologica Italiana*, 44: 35-45.

Girone A., Capotondi L., Ciaranfi N., Di Leo P., Lirer F., Maiorano P., Marino M., Pelosi N. & Pulice I. (2013a). Paleoenvironmental changes at the lower Pleistocene Montalbano Jonico section (southern Italy): global versus regional signals. *Palaeogeography, Palaeoclimatology, Palaeoecology*, 371: 62-79.

Girone A., Maiorano P., Marino M. & Kucera M. (2013b). Calcareous plankton response to orbital and millennial-scale climate changes across the Middle Pleistocene in the western Mediterranean. *Palaeogeography, Palaeoclimatology, Palaeoecology*, 392: 105-116.

Gliozzi E., Abbazzi L., Argenti P., Azzaroli A., Caloi L., Capasso Barbato L., Di Stefano G., Esu D., Ficcarelli G., Girotti O., Kotsakis T., Masini F., Mazza P., Mezzabotta C., Palombo M.R., Petronio C., Rook L., Sala B., Sardella R., Zanalda E. & Torre D. (1997). Biochronology of selected mammals, mollusks and ostracods from the Middle Pliocene to the Late Pleistocene in Italy. The state of the art. *Rivista Italiana di Paleontologia e Stratigrafia*, 103: 368-388.

Gradstein F.M. & Ogg J.G. (2020). The chronostratigraphic scale. In Gradstein F., Ogg J.G., Schmitz M.D. & Ogg G.M. (eds), Geologic time scale 2020, Elsevier: 21-32.

Gradstein F.M., Ogg J.G. & Smith A.G. (2004). Construction and summary of geologic time scale. In Gradstein F.M., Ogg J.G. & Smith A. (eds), A Geologic Time Scale. Cambridge University Press: 455-464.

Grant K.M., Amarathunga U., Amies J.D., Hu P., Qian Y., Penny T., Rodriguez-Sanz L., Zhao X., Heslop D., Liebrand D., Hennekam R., Westerhold T., Gilmore S., Lourens L.J., Roberts A.P. & Rohling E.J. (2022). Organic carbon burial in Mediterranean sapropels intensified during Green Sahara Periods since 3.2 Myr ago. *Communications Earth and Environment*, 3: 11.

Hay W.W., Mohler H.P., Roth P.H., Schmidt R.R. & Boudreaux J.E. (1967). Calcareous nannoplankton zonation of the Cenozoic of the Gulf Coast and Caribbean-Antillean area, and transoceanic correlation. *Transactions of the Gulf-Coast Association of Geological Societies*, 17: 428-480.

Haywood A.M. & Valdes P.J. (2004). Modelling Middle Pliocene warmth: contribution of atmosphere, oceans and cryosphere. *Earth Planetary Science Letters*, 218: 363-377.

Head M.J. (2021). Review of the Early-Middle Pleistocene boundary and Marine Isotope Stage 19. *Progress in Earth and Planetary Science*, 8: 50.

Head M.J. & Gibbard P.J. (2015). Early-Middle Pleistocene transitions: linking terrestrial and marine realms. *Quaternary International*, 389: 7-46.

Head M.J., Gibbard P. & Salvador A. (2008). The Quaternary: its character and definition. *Episodes*, 31: 234-238.

Hemleben C., Spindler M. & Anderson O.R. (1989). Taxonomy and species features. Modern Planktonic Foraminifera. Springer New York: 8-32.

Herbert T.D., Gideon N.G. & Cleaveland Peterson L. (2015). Evolution of Mediterranean Sea surface temperatures 3.5-1.5 Ma: regional and hemispheric influences. *Earth Planetary Science Letters*, 409: 307-318.

Herbert T.D., Lawrence K.T., Tzanova A., Peterson L.C., Caballero-Gill R.P. & Kelly C.S. (2018). (Table S2) SST estimates as a function of age, Mediterranean sites. PANGAEA. https://doi.org/10.1594/PANGAEA.885603.

Hilgen F.J. (1990). Closing the gap in the Plio-Pleistocene boundary stratotype sequence of Crotone (southern Italy). *Newsletters on Stratigraphy*, 22: 43-51.

Hilgen F.J. (1991). Astronomical calibration of Gauss to Matuyama sapropels in the Mediterranean and implication for the geomagnetic polarity time scale. *Earth and Planetary Science Letters*, 104: 226-244.

Howell M.W., Thunell R., Tappa E., Rio D. & Sprovieri R. (1988). Late Neogene laminated and opal-rich facies from the Mediterranean region: geochemical evidence for mechanisms of formation. *Palaeogeography, Palaeoclimatology, Palaeoecology*, 64: 265-286.

Joannin S., Quillévéré F., Suc J.P., Lécuyer C. & Martineau F. (2007). Early Pleistocene climate changes in the central Mediterranean region as inferred from integrated pollen and foraminiferal stable isotope analyses. *Quaternary Research*, 67: 264-274.

Joannin S., Ciaranfi N. & Stefanelli S. (2008). Vegetation changes during the late early Pleistocene at Montalbano Jonico (Province of Matera, southern Italy) based on pollen analysis. *Palaeogeography, Palaeoclimatology, Palaeoecology*, 270: 92-101.
Kaboth-Bahr S., Bahr A., Stepanek C., Catunda M.C.A., Karas C., Ziegler M., García-Gallardo A. & Grunert P. (2021). Mediterranean heat injection to the North Atlantic delayed the intensification of Northern Hemisphere glaciations. *Communications Earth & Environment*, 2: 158.

Kamptner E. (1943). Zur Revision der Coccolithineen-Spezies *Pontosphaera huxleyi* Lohm. *Anzeiger der (Kaiserlichen) Akademie der Wissenschaften, Wien*, 80: 43-49.

Kamptner E. (1963). Coccolithineen-Skelettreste aus Tiefseeablagerungen des Pazifischen Ozeans. *Annalen des Naturhistorischen Museums in Wien*, 66: 139-204.

Kandiano E.S., Bauch H.A., Fahl K., Helmke J.P., Röhl U., Pérez-Folgado M. & Cacho I. (2012). The meridional temperature gradient in the eastern North Atlantic during MIS11 and its link to the ocean-atmosphere system. *Palaeogeography, Palaeoclimatology, Palaeoecology*, 333-334: 24-39.

Kindler P. & Hearty P.J. (2000). Elevated marine terraces from Eleuthera (Bahamas) and Bermuda: sedimentological, petrographic, and geochronological evidence for important deglaciation events during the middle Pleistocene. *Global and Planetary Change*, 24: 41-58.

Klotz S., Fauquette S., Combourieu-Nebout N., Uhl D., Suc J.P. & Mosbrugger V. (2006). Seasonality intensification and long-term winter cooling as a part of the Late Pliocene climate development. *Earth Planetary Science Letters*, 241: 174-187.

Kroon D., Alexander I., Little M., Lourens L., Matthewson A., Robertson A. & Sakamoto T. (1998). Oxygen isotope and sapropel stratigraphy in the eastern Mediterranean during the last 3.2 million years. *Proceedings of the Ocean Drilling Program, Scientific Results*, 160: 181-189.

Kulhanek D.K. (2009). Calcareous nannoplankton as paleoceanographic and biostratigraphic proxies: Examples from the mid-Cretaceous equatorial Atlantic (ODP leg 207) and Pleistocene of the Antarctic Peninsula (NBP0602A) and North Atlantic (IODP Exp. 306). 210 pp. Ph.D. Dissertation, College of Arts and Sciences, Florida State University.

Lang N. & Wolff E.W. (2011). Interglacial and glacial variability from the last 800 ka in marine, ice and terrestrial archives. *Climate of the Past*, 7: 361-380.

Larrasoña J.C., Roberts A.P., Rohling E.J., Winklhofer M. & Wehausen R. (2003). Three million years of monsoon variability over the northern Sahara. *Climate Dynamics*, 21: 689-698.

Leroy S. (2007). Progress in palynology of the Gelasian-Calabrian stages in Europe: ten messages. *Revue de Micropaléontologie*, 50: 293-308.

Linnaeus C. (1753). Species Plantarum. Exhibentes plantas rite cognitas ad genera relates. 560 pp. Laurentii Salvii, Stockholm.

Lionello P., Giorgi F. & Rohling E. (2023). Mediterranean climate: Past, present and future. *In* Schroeder K. & Chiggiato J. (eds), *Oceanography of the Mediterranean Sea*. Elsevier, Amsterdam: 41-91.

Lirer F., Foresi L.M., Iaccarino S.M., Salvatorini G., Turco E., Cosentino C., Sierto F.J. & Caruso A. (2019). Mediterranean Neogene planktonic foraminifer biozonation and biochronology. *Earth-Science Reviews*, 196: 102869.

Lisiecki L.E. & Raymo M.E. (2005). A Pliocene-Pleistocene stack of 57 globally distributed benthic $\delta^{18}$O records. *Paleoceanography*, 20: PA1003.

Lona F. & Bertoldi R. (1972). La storia del Plio-Pleistocene italiano in alcune sequenze vegetazionali lacustri e marine. *Atti della Accademia Nazionale dei Lincei*, 11: 1-45.

Lourens L.J. (2004). Revised tuning of Ocean Drilling Program Site 964 and KC01B (Mediterranean) and implications for the $\delta^{18}$O, tephra, calcareous nannofossil, and geomagnetic reversal chronologies of the past 1.1 Myr. *Paleoceanography*, 19: PA3010.

Lourens L.J., Hilgen F.J., Gudjonsson L. & Zachariasse W.J. (1992). Late Pliocene to early Pleistocene astronomically forced sea surface productivity and temperature variations in the Mediterranean. *Marine Micropaleontology*, 19: 49-78.

Lourens L.J., Antonarakou F., Hilgen F.J., Van Hoof A.M., Vergnaud-Grazzini C. & Zacharias W.J. (1996a). Evaluation of the Plio-Pleistocene astronomical timescale. *Paleoceanography*, 11: 491-493.

Lourens L.J., Hilgen F.J., Raffi I. & Vergnaud-Grazzini C. (1996b). Early Pleistocene chronology of the Vrica section (Calabria, Italy). *Paleoceanography*, 11: 797-812.

Lourens L.J., Hilgen F.J. & Raffi I. (1998). Base of large *Gephyrocapsa* and astronomical calibration of early Pleistocene sapropels in Site 967 and Hole 969D: solving the chronology of the Vrica section (Calabria, Italy). *Proceedings of the Ocean Drilling Program, Scientific Results*, 160: 191-198.

Lourens L.J., Hilgen F.J., Laskar J., Shackleton N.J. & Wilson D. (2004). The Neogene Period. *In* Gradstein F.M., Ogg J.G. & Smith A.G. (eds), *A Geologic Time Scale 2004*. Cambridge University Press: 409-440.

Magri D., Di Rita F., Aranbarri J., Fletcher W. & González-Sampériz P. (2017). Quaternary disappearance of tree taxa from Southern Europe: Timing and trends. *Quaternary Science Reviews*, 163: 23-55.

Maiorano P., Marino M., Di Stefano E. & Ciaranfi N. (2004). Calcareous nannofossil events in the Lower-Middle Pleistocene transition at the Montalbano Jonico section and ODP Site 964: calibration with isotope and sapropel stratigraphy. *Rivista Italiana di Paleontologia e Stratigrafia*, 110: 547-557.

Maiorano P., Aiello G., Barra D., Di Leo P., Joannin S., Lirer F., Marino M., Pappalardo A., Capotondi L., Ciaranfi N. & Stefanelli S. (2008). Paleoenvironmental changes during sapropel 19 (i-cycle 90) deposition: evidences from geochemical, mineralogical and micropaleontological proxies in the mid Pleistocene Montalbano Jonico land section (southern Italy). *Palaeogeography, Palaeoclimatology, Palaeoecology*, 257: 308-334.

Maiorano P., Capotondi L., Ciaranfi N., Girone A., Lirer F., Marino M., Pelosi N., Petrosino P. & Piscitelli A. (2010). Vrica-Crotone and Montalbano Jonico sections: a potential unit-stratotype of the Calabrian Stage. *Episodes*, 33: 218-233.

Maiorano P., Tarantino F., Marino M. & Girone A. (2013a). A paleoecological and paleobiogeographic evaluation of *Helicosphaera inversa* (Gartner) Theodoridis and the diachrony of its first occurrence. *Marine Micropaleontology*, 104: 14-24.

Maiorano P., Tarantino F., Marino M. & De Lange G.J. (2013b). Paleoenvironmental conditions at Core KC01B (Ionian Sea) through MIS 13-9: Evidence from calcareous nannofossil assemblages. *Quaternary International*, 288: 97-111.

Maiorano P., Marino M., Balestra B., Flores J.A., Hodell D.A. & Rodrigues T. (2015). Coccolithophore variability from the Shackleton Site (IODP Site U1385) through MIS 16-10. *Global and Planetary Change*, 133: 35-48.

Maiorano P., Bertini A., Capolongo D., Eramo G., Gallicchio S., Girone A., Pinto D., Toti F., Ventruti G. & Marino M. (2016a). Climate signatures through the Marine Isotope Stage 19 in the Montalbano Jonico section (Southern Italy): a land-sea perspective. *Palaeogeography, Palaeoclimatology, Palaeoecology*, 461: 341-361.

Maiorano P., Girone A., Marino M., Kucera M. & Pelosi N. (2016b). Sea surface water variability during the Mid-Brunhes inferred from calcareous plankton in the western Mediterranean (ODP Site 975). *Palaeogeography, Palaeoclimatology, Palaeoecology*, 459: 229-248.

Maiorano P., Herbert T.D., Marino M., Bassinot F., Bazzicalupo P., Bertini A., Girone A., Nomade S. & Ciaranfi N. (2021). Paleoproductivity modes in central Mediterranean during MIS 20-MIS 18: calcareous plankton and alkenone variability. *Paleoceanography and Paleoclimatology*, 36: e2021PA004259.

Maldonado A., Bohoyo F., Galindo-Zaldivar J., Hernandez-Molina F.J., Lobo F.J., Lodolo E., Martos Y.M., Pérez L.F., Schreider A.A. & Somoza L. (2014). A model of oceanic development by ridge jumping: Opening of the Scotia Sea. *Global and Planetary Change*, 123: 152-173.
Marino M., Maiorano P., Tarantino F., Voelker A., Capotondi L., Girone A., Lirer F., Flores J.-A. & Naafs B.D.A. (2014). Coccolithophores as proxy of seawater changes at orbital-to-millennial scale during middle Pleistocene Marine Isotope Stages 14-9 in North Atlantic core MD01-2446. *Paleoceanography and Paleoclimatology*, 29: 518-532.

Marino M., Bertini A., Ciaranfi N., Aiello G., Barra D., Gallicchio S., Girone A., LaPerna R., Lirer F., Maiorano P., Petrosino P. & Toti F. (2015). Paleoenvironmental and climatostratigraphic insights for Marine Isotope Stage 19 (Pleistocene) at the Montalbano Jonico succession, south Italy. *Quaternary International*, 383: 104-115.

Marino M., Girone A., Maiorano P., Di Renzo R., Piscitelli A. & Flores J.A. (2018). Calcareous plankton 1141 and the mid-Brunhes climate variability in the Alboran Sea (ODP Site 977). *Palaeogeography, Palaeoclimatology, Palaeoecology*, 508: 91-106.

Marino M., Girone A., Gallicchio S., Herbert T., Addante M., Bazzicalupo P., Quivelli O., Bassinot F., Bertini A., Nomade S., Ciaranfi N. & Maiorano P. (2020). Climate variability during MIS 20-18 as recorded by alkenone-SST and calcareous plankton in the Ionian Basin (central Mediterranean). *Palaeogeography, Palaeoclimatology, Palaeoecology*, 560: 110027.

Marino M., Maiorano P., Girone A., Gallicchio S., Bassinot F., Nomade S., Simon Q., Bertini A., Herbert T., Petrosino P., Addante M., La Perna R., Trotta S. & Ciaranfi N. (2024). The Ideale section at Montalbano Jonico (southern Italy): a Standard Auxiliary Boundary Stratotype (SABS) for the Middle Pleistocene Subseries of the Quaternary System. *Episodes*, 47: 121-146.

Martinetto E., Bertini A., Basilici G., Baldanza A., Bizzarri R., Cherin M., Gentili S. & Pontini M.R. (2014). The plant record of the Dunarobba and Pietrafitta sites in the Plio-Pleistocene palaeoenvironmental context of central Italy. *Alpine Mediterranean Quaternary*, 27: 29-72.

Martrat B., Grimalt J.O., Lopez-Martinez C., Cacho I., Sierro F.J., Flores J.A., Zahn R., Canals M., Curtis J.H. & Hodell D.A. (2004). Abrupt temperature changes in the Western Mediterranean over the past 250,000 years. *Science*, 306: 1762-1765.

Masson-Delmotte V., Stenni B., Pol K., Braconnot P., Cattani O., Falourd S., Kageyama M., Jouzel J., Landais A., Minster B., Barnola J.M., Chappellaz J., Krinner G., Johnsen S., Röthlisberger R., Hansen J., Mikolajewicz U. & Otto-Bliesner B. (2010). EPICA Dome C record of glacial and interglacial intensities. *Quaternary Science Reviews*, 29: 113-128.

McClymont E.L., Sosdian S.M., Rosell-Melé A. & Rosenthal Y. (2013). Pleistocene sea-surface temperature evolution: Early cooling, delayed glacial intensification, and implications for the mid-Pleistocene climate transition. *Earth-Science Reviews*, 123: 173-193.

McClymont E.L., Ho S.L., Ford H.L., Bailey I., Berke M.A., Bolton C.T., De Schepper S., Grant G.R., Groeneveld J., Inglis G.N., Karas C., Patterson M.O., Swann G.E.A., Thirumalai K., White S.M., Alonso-Garcia M., Anand P., Hoogakker B.A.A., Littler K., Petrick B.F., Risebrobakken B., Abell J.T., Crocker A.J., de Graaf F., Feakins S.J., Hargreaves J.C., Jones C.L., Markowska M., Ratnayake A.S., Stepanek C. & Tangunan D. (2023). Climate evolution through the onset and intensification of Northern Hemisphere Glaciation. *Reviews of Geophysics*, 61: e2022RG000793.

Milker Y., Rachmayani R., Weinkauf M.F.G., Prange M., Raitzsch M., Schulz M. & Kucera M. (2013). Global and regional sea surface temperature trends during Marine Isotope Stage 11. *Climate of the Past*, 9: 2231-2252.

Mondanaro A., Dominici S. & Danise S. (2024). Response of Mediterranean Sea bivalves to Pliocene-Pleistocene environmental changes. *Palaeontology*, 67: e12696.

Moscariello A., Ravazzi C., Brauer A., Mangili C., Chiesa S., Rossi S., de Beaulieu J.L. & Reille M. (2000). A long lacustrine

record from the Piânico-Sèllere basin (Middle-Late Pleistocene, northern Italy). *Quaternary International*, 73: 47-68.

Mudelsee M. & Raymo M.E. (2005). Slow dynamics of the Northern Hemisphere glaciation. *Paleoceanography*, 20: PA4022.

Mudelsee M. & Schulz M. (1997). The Mid-Pleistocene climate transition: onset of 100 ka cycle lags ice volume build up by 280 ka. *Earth and Planetary Science Letters*, 151: 117-123.

Muttoni G., Ravazzi C., Breda M., Pini R., Laj C., Kissel C., Mazaud A. & Garzanti E. (2007). Magnetostratigraphic dating of an intensification of glacial activity in the southern Italian Alps during Marine Isotope Stage 22. *Quaternary Research*, 67: 161-173.

Naafs B.D.A., Hefter J., Ferretti P., Stein R. & Haug G.H. (2011). Sea surface temperatures did not control the first occurrence of Hudson Strait Heinrich events during MIS 16. *Paleoceanography*, 26: PA4201.

Naafs B.D.A., Hefter J. & Stein R. (2013). Millennial-scale ice rafting events and Hudson Strait Heinrich(-like) events during the late Pliocene and Pleistocene: a review. *Quaternary Science Review*, 80: 1-28.

Naafs B.D.A., Voelker A.H.L., Karas C., Andersen N. & Sierro F.J. (2020). Repeated Near-Collapse of the Pliocene Sea Surface Temperature Gradient in the North Atlantic. *Paleoceanography and Paleoclimatology*, 35: e2020PA003905.

Nakagawa H., Niitsuma N., Takayama T., Matoba Y., Oda M., Tokunaga S., Kitazato H., Sakai T. & Koizumi I. (1997). The magnetostratigraphy of the Vrica section, Italy, and its correlation with the Plio-Pleistocene of the Boso Penin, Japan. *In Van Couvering J.A.* (ed.), The Pleistocene Boundary and the Beginning of the Quaternary. Cambridge University Press, Cambridge: 46-56.

Natland M.L. (1938). New Species of Foraminifera from off the West Coast of North America and from the Later Tertiary of the Los Angeles Basin. *Bulletin of the Scripps Institute of Oceanography, Technical Series*, 4: 137-164.

Nomade S., Bassinot F., Marino M., Simon Q., Dewilde F., Maiorano P., Isguder G., Blamart D., Girone A., Scao V., Pereira A., Toti F., Bertini A., Combourieu-Nebout N., Peral M., Bourlès D.L., Petrosino P., Gallicchio S. & Ciaranfi N. (2019). High-resolution foraminifer stable isotope record of MIS 19 at Montalbano Jonico, southern Italy: A window into Mediterranean climatic variability during a low-eccentricity interglacial. *Quaternary Science Reviews*, 205: 106-125.

Pagani M., Caldeira K., Berner R. & Beerling D.J. (2009). The role of terrestrial plants in limiting atmospheric CO₂ decline over the past 24 million years. *Nature*, 460: 85-88.

Palombo M.R. (2004). Biochronology of the Plio-Pleistocene mammalian faunas of Italian peninsula: knowledge, problems and perspectives. *Il Quaternario*, 17: 565-582.

Palombo M.R. (2007). Large mammal guilds on the Italian peninsula from the Pliocene to the Late Pleistocene: new methodological approaches. *Courier Forschungsinstitut Senckenberg*, 259: 13-46.

Parker F.L. (1958). Eastern Mediterranean foraminifera. *Reports of the Swedish Deep-Sea Expedition*, 1947-1948, 8: 217-283.

Pasini G. & Colalongo M.L. (1994). Proposal for the erection of the Santerrian/Emilian boundary-stratotype (lower Pleistocene) and new data on the Pliocene/Pleistocene boundary stratotype. *Bollettino della Società Paleontologica Italiana*, 33: 101-120.

Pasini G. & Colalongo M.L. (1997). The Pliocene-Pleistocene boundary stratotype at Vrica, Italy. *In Van Couvering J.A.* (ed.), The Pleistocene Boundary and the Beginning of the Quaternary. Cambridge University Press, Cambridge: 15-45.

Past Interglacials Working Group of PAGES (2016). Interglacials of the last 800,000 years. *Reviews of Geophysics*, 54: 162-219.

Petrick B., McClymont E.L., Littler K., Rosell-Melé A., Clarkson M.O., Maslin M., Röhl U., Shevenell A.E. & Pancost R.D. (2018). Oceanographic and climatic evolution of the southeastern subtropical Atlantic over the last 3.5 Ma. *Earth and Planetary Science Letters*, 492: 12-21.
Petrosino P., Jicha B.R., Mazzeo F.C., Ciaranfi N., Girone A., Maiorano P. & Marino M. (2015). The Montalbano Jonico marine succession: An archive for distal tephra layers at the Early-Middle Pleistocene boundary in southern Italy. *Quaternary International*, 383: 89-103.

Pini R., Bertini A., Martinetto E. & Vassio E. (2014). The Pleistocene flora of northern Italy. Palaeobotany of Italy. *Publication of the Museum of Nature South Tyrol*, 9: 290-307.

Pisias N.G. & Moore Jr. T.C. (1981). The evolution of the Pleistocene climate: a time series approach *Earth and Planetary Science Letters*, 52: 450-458.

Pontini M.R. (1997). Analisi palinologica e interpretazione paleoclimatica dei depositi continentali del Bacino Tiberino (Umbria, Italia). 134 pp. Unpublished Ph.D. Thesis, Università di Perugia, Italy.

Pross J., Christianis K., Fischer T., Fletcher W.J., Hardiman M., Kalaitzidis S., Knipping M., Kotthoff U., Milner A.M., Muller U.C., Schmiedl G., Siavalas G., Tzedakis P.C. & Wulf S. (2015). The 1.35-Ma-long terrestrial climate archive of Tenaghi Philippon, northeastern Greece: Evolution, exploration, and perspectives for future research. *Newsletters on Stratigraphy*, 48: 253-276.

Quivelli O., Marino M., Rodrigues T., Girone A., Maiorano P., Abrantes F., Salgueiro E. & Bassinot F. (2020). Surface and deep-water variability in the Western Mediterranean (ODP Site 975) during insolation cycle 74: high resolution calcareous plankton and molecular biomarker signals. *Palaeogeography, Palaeoclimatology, Palaeoecology*, 542: 109583.

Quivelli O., Marino M., Rodrigues T., Girone A., Maiorano P., Bertini A., Niccolini G., Trotta S. & Bassinot F. (2021). Multiproxy record of suborbital-scale climate changes in the Algero-Balearic Basin during late MIS 20 - Termination IX. *Quaternary Science Review*, 260: 106916.

Raffi I., Backman J., Fornaciari E., Pálike H., Rio D., Lourenς L. & Hilgen F. (2006). A review of calcareous nannofossil astrobiochronology encompassing the past 25 million years. *Quaternary Science Review*, 25: 3113-3137.

Ravazzi C. (2003). Gli Antichi Bacini Lacustri e i Fossili di Leffe, Ranica e Piánico-Sellere. 176 pp. Quaderni di Geodinamica Alpina e Quaternaria, CNR Special Publications, Bergamo, Italy.

Ravazzi C. & Rossignol-Strick M. (1995). Vegetation change in a climatic cycle of early Pleistocene age in the Leffe basin (Northern Italy). *Palaeogeography, Palaeoclimatology, Palaeoecology*, 117: 105-122.

Ravazzi C., Pini R., Breda M., Martinetto E., Muttoni G., Chiesa S., Confortini F. & Egli R. (2005). The lacustrine deposits of Fornaci di Ranica (late Early Pleistocene, Italian Pre-Alps): stratigraphy, palaeoenvironment and geological evolution. *Quaternary International*, 131: 35-58.

Ravelo A.C., Andreasen D.H., Lyle M., Olivarez Lyle A. & Wara M.W. (2004). Regional climate shifts caused by gradual global cooling in the Pliocene epoch. *Nature*, 429: 263-267.

Regattieri E., Giaccio B., Galli P., Nomade S., Peronace E., Messina P., Sposato A., Boschi C. & Gemelli M. (2015). A multi-proxy record of MIS 11-12 deglaciation and glacial MIS 12 instability from the Sulmona Basin (central Italy). *Quaternary Science Review*, 32: 129-145.

Regattieri E., Giaccio B., Mannella G., Zanchetta G., Nomade S., Tognarelli A., Perchiazzi N., Vogel H., Boschi C., Neil R.D., Wagner B., Gemelli M. & Tzedakis P. (2019). Frequency and dynamics of millennial-scale variability during Marine Isotope Stage 19: Insights from the Sulmona Basin (central Italy). *Quaternary Science Reviews*, 214: 28-43.

Rio D., Raffi I. & Villa G. (1990). Pliocene-Pleistocene distribution patterns in the Western Mediterranean. *Proceedings of the Ocean Drilling Program, Scientific Results*, 107: 513-533.

Rio D., Backman J. & Raffi I. (1996). Calcareous nannofossil Biochronology and the Pliocene-Pleistocene Boundary. *In Van Couvering J.A. (ed.)*, The Pleistocene Boundary and the

Beginning of the Quaternary. Cambridge University Press, Cambridge: 63-78.

Rio D., Channell J.E.T., Bertoldi R., Poli M.S., Vergerio P.P., Raffi I., Sprovieri R. & Thunell R.C. (1997). Pliocene sapropels in the northern Adriatic area: chronology and paleoenvironmental significance. *Palaeogeography, Palaeoclimatology, Palaeoecology*, 135: 1-25.

Rio D., Sprovieri R., Castradori D. & Di Stefano E. (1998). The Gelasian Stage (Upper Pliocene): a new unit of the global standard chronostratigraphic scale. *Episodes*, 21: 82-87.

Rodrigues T., Voelker A.H.L., Grimalt J.O., Abrantes F. & Naughton F. (2011). Iberian Margin sea surface temperature during MIS 15 to 9 (580-300 ka): Glacial suborbital variability versus interglacial 1310 stability. *Paleoceanography*, 26: PA1204.

Rodrigues T., Alonso-García M., Hodell D.A., Rufino M., Naughton F., Grimalt J.O., Voelker A.H.L. & Abrantes F. (2017). A 1-Ma record of sea surface temperature and extreme cooling events in the North 1313 Atlantic: A perspective from the Iberian Margin. *Quaternary Science Review*, 172: 118-130.

Rögl F. & Bolli H. M. (1973). Holocene to Pleistocene planktonic foraminifera of LEG 15, site 147 (Cariaco Basin (Trench), Caribbean Sea) and their climatic interpretation. *Initial Reports of the Deep Sea Drilling Project*, 15: 553-579.

Rohling E.J., Marino G. & Grant K.M. (2015). Mediterranean climate and oceanography, and the periodic development of anoxic events (sapropels). *Earth-Science Reviews*, 143: 62-97.

Rook L. & Martínez-Navarro B. (2010). Villafranchian: The long story of a Plio-Pleistocene European large mammal biochronologic unit. *Quaternary International*, 219: 134-144.

Rossignol M. (1962). Analyse pollinique de sédiments marins quaternaires en Israël II. - Sédiments pléistocènes. *Pollen et Spores*, 4: 121-148.

Rossignol M. (1964). Hystrichosphères du Quaternaire en Méditerranée orientale, dans les sédiments Pléistocènes et les boues marines actuelles. *Revue de micropaléontologie*, 7: 83-99.

Ruddiman W.F., Raymo M.E., Martinson D.G., Clement B.M. & Backman J. (1989). Pleistocene evolution: northern hemisphere ice sheets and North Atlantic Ocean. *Paleoceanography*, 4: 353-412.

Ruggieri E., Herbert T., Lawrence K.T. & Lawrence C.E. (2009). Change point method for detecting regime shifts in paleoclimatic time series: application to $\delta^{18}O$ time series of the Plio-Pleistocene. *Paleoceanography*, 24: PA1204.

Sabato L., Bertini A., Masini F., Albianelli A., Napoleone G. & Pieri P. (2005). The lower and middle Pleistocene geological record of the San Lorenzo lacustrine sequence in Sant'Arcangelo basin. (Southern Apennines, Italy). *Quaternary International*, 131: 59-69.

Sadori L., Bertini A., Combourieu-Nebout N., Kouli K., Mariotti Lippi M., Roberts N. & Mercuri A.M. (2013). Palynology and Mediterranean vegetation history. *Flora Mediterranea*, 23: 141-156.

Sánchez-Goñi M.F., Rodrigues T., Hodell D.A., Polanco-Martínez J.M., Alonso-García M., Hernández-Almeida I., Desprat S. & Ferretti P. (2016). Tropically-driven climate shifts in southwestern Europe during MIS 19, a low eccentricity interglacial. *Earth Planetary Science Letters*, 448: 81-93.

Sánchez-Goñi M. F., Extier T., Polanco-Martínez J. M., Zorzi C., Rodrigues T. & Bahr A. (2023). Moist and warm conditions in Eurasia during the last glacial of the Middle Pleistocene Transition. *Nature Communications*, 14: 2700.

Sarjeant W.A.S. (1970). The genus *Spiniferites* Mantell, 1850 (Dinophyceae). *Grana*, 10: 74-78.

Sato T. & Takayama T. (1992). A stratigraphically significant new species of the calcareous nannofossil *Reticulofenestra asanoi*. In Ishizaki K. & Sato T. (eds), Century of Japanese Micropalaeontology. Terra Scientific Tokyo: 457-460.

Scherer R.P., Bohaty S.M., Dunbar R.B., Esper O., Flores J.A., Gersonde R., Harwood D.M., Roberts A.P. & Taviani M.
(2008). Antarctic records of precession-paced insolation-driven warming during early Pleistocene Marine Isotope Stage 31. *Geophysical Research Letters*, 35: L03505.

Schröter J.S. (1783). Einleitung in die Conchylienkenntniss nach Linné. 860 pp. Johan Jacob Gebauer, Halle.

Schueth J. & Bralower T.J. (2015). The relationship between environmental change and the extinction of the nannoplankton *Discoaster* in the early Pleistocene. *Paleoceanography*, 30: 863-876.

Selli R. (1967). The Plio-Pleistocene boundary in Italian marine sections and its relationship to continental stratigraphies. *Progress in Oceanography*, 4: 67-87.

Serrano F. (2020). An approach to the paleoceanographic characteristics of the sea-surface at the western Mediterranean (Balearic area) during the Pliocene and Gelasian. *Geosciences*, 10: 1-21.

Shackleton N.J. & Opdyke N.D. (1976). Oxygen-isotope and paleomagnetic stratigraphy of Pacific core V28-239: late Pliocene to latest Pleistocene. *Geological Society of America Memoirs*, 145: 449-464.

Shackleton N.J., Hagelberg T.K. & Crowhurst S.J. (1995). Evaluating the success of astronomical tuning: pitfalls of using coherence as a criterion for assessing pre-Pleistocene timescales. *Paleoceanography*, 10: 693-697.

Simon Q., Bourlès L.D., Bassinot F., Nomade S., Marino M., Ciaranfi N., Girone A., Maiorano P., Thouveny N., Choya S., Dewil F., Scao V., Isguder G., Blamart D. & ASTER Team (2017). Authigenic $^{10}$Be/$^{9}$Be ratio signature of the Matuyama-Brunhes boundary in the Montalbano Jonico marine succession. *Earth and Planetary Science Letters*, 460: 255-267.

Spaak P. (1983). Accuracy in correlation and ecological aspects of the planktonic foraminiferal zonation of the Mediterranean Pliocene. *Utrecht Micropaleontological Bulletin*, 28: 1-159.

Sprovieri R., Di Stefano E., Incarbona A. & Oppo D.W. (2006) Suborbital climate variability during Marine Isotopic Stage 5 in the central Mediterranean basin: evidence from calcareous plankton record. *Quaternary Sciences Review*, 25: 2332-2342.

Stefanelli S. (2003). Benthic foraminiferal assemblages as tools for paleo-environmental reconstruction of the early-middle Pleistocene Montalbano Jonico composite section. *Bollettino della Società Paleontologica Italiana*, 42: 281-299.

Stefanelli S. (2004). Cyclic stages in oxygenation based on foraminiferal microhabitats: early-middle Pleistocene, Lucania basin, southern Italy. *Journal of Micropaleontology*, 23: 81-95.

Stefanelli S., Capotondi L. & Ciaranfi N. (2005). Foraminiferal record and environmental changes during the deposition of early-middle Pleistocene sapropels in southern Italy. *Palaeogeography, Palaeoclimatology, Palaeoecology*, 216: 27-52.

Stein R., Hefter J., Grützner J., Voelker A. & Naafs B.D.A. (2009). Variability of surface water characteristics and Heinrich-like events in the Pleistocene mid latitude North Atlantic Ocean: Biomarker 1377 and XRD records from IODP Site U1313 (MIS 16-9). *Paleoceanography*, 24: PA2203.

Suc J.P., Bertini A., Combourieu-Nebout N., Diniz F., Leroy S., Russo Ermolli E., Zheng Z., Bessais E. & Ferrier J. (1995). Structure of west Mediterranean and climate since 5.3 Ma. *Acta Zoologica Cracoviensa*, 38: 3-16.

Suc J.P., Combourieu-Nebout N., Seret G., Popescu S.M., Klotz S., Gautier F., Clauzon G., Westgate J., Insinga D. & Sandhu A.S. (2010). The Crotone series: a synthesis and new data. *Quaternary International*, 219: 121-133.

Suc J.P., Popescu S.M., Fauquette S., Bessedik M., Jiménez-Moreno G., Bachiri Taoufiq N., Zheng Z., Medial F. & Klotz S. (2018). Reconstruction of Mediterranean flora, vegetation and climate for the last 23 million years based on an extensive pollen dataset. *Ecologia Mediterranea*, 44: 53-85.

Suganuma Y., Okada M., Head M.J., Kameo K., Haneda Y., Hayashi H., Irizuki T., Itaki T., Izumi K., Kubota Y., Nakazato H., Nishida N., Okuda M., Satoguchi Y., Simon Q., Takeshita Y. & the

Chiba composite section community members (2021). Formal ratification of the Global Boundary Stratotype Section and Point (GSSP) for the Chibanian Stage and Middle Pleistocene Subseries of the Quaternary System: the Chiba Section, Japan. *Episodes*, 44: 317-347.

Tan S.H. (1927). *Discoasteridae* incertae sedis. *Proceedings of the Koninklijke Nederlandse Akademie van Wetenschappen. Section of Sciences*, 30: 411-419.

Tangunan D.N., Baumann K.H., Just J., LeVay L.J., Barker S., Brentegani L., De Vleeschouwer D., Hall I.R., Hemming S. & Norris R. (2018). The last 1 million years of the extinct genus *Discoaster*: Plio-Pleistocene environment and productivity at Site U1476 (Mozambique Channel). *Palaeogeography, Palaeoclimatology, Palaeoecology*, 505: 187-197.

Toti F. (2015). Interglacial vegetation patterns at the Early-Middle Pleistocene transition: a point of view from Montalbano Jonico section (southern Italy). *Alpine and Mediterranean Quaternary*, 28: 131-143.

Toti F. & Bertini A. (2018). Mediterranean pollen stratigraphy across the Early-Middle Pleistocene boundary. *Alpine and Mediterranean Quaternary*, 31: 199-203.

Toti F., Bertini A., Girone A., Marino M., Maiorano P., Bassinot F., Combourieu-Nebout N., Nomade S. & Buccianti A. (2020). Marine and terrestrial climate variability in the western Mediterranean Sea during marine isotope stages 20 and 19. *Quaternary Science Reviews*, 243: 106486.

Trotta S., Marino M., Maiorano P. & Girone A. (2019). Climate variability through MIS 20-MIS 19 in core KC01B, Ionian Basin (central Mediterranean Sea). *Alpine and Mediterranean Quaternary*, 32: 1-15.

Tzedakis P.C. (2005). Towards an understanding of the response of southern European vegetation to orbital and suborbital climate variability. *Quaternary Sciences Review*, 25: 1585-1599.

Tzedakis P.C. (2007). Seven ambiguities in the Mediterranean palaeoenvironmental narrative. *Quaternary Sciences Review*, 26: 2042-2066.

Tzedakis P.C., Channell J.E.T., Hodell D.A., Kleiven H.F. & Skinner L.C. (2012). Determining the natural length of the current interglacial. *Nature Geoscience*, 5: 138-141.

Tzedakis P.C., Crucifix M., Mitsui T. & Wolff E.W. (2017). A simple rule to determine which insolation cycles lead to interglacials. *Nature*, 542(7642): 427-432.

Vannacci M. (2016). Cambiamenti climatici durante lo stadio isotopico 22 come registrato nella successione marina di Montalbano Jonico (Italia meridionale) tramite indagine palinologica. 70 pp. Unpublished Master thesis, Università di Firenze.

Voelker A.H.L., Rodrigues T., Billups K., Oppo D., McManus J., Stein R., Hefter J. & Grimalt J.O. (2010). Variations in mid-latitude North Atlantic surface water properties during the mid-Brunhes (MIS 9-14) and their implications for the thermohaline circulation. *Climate of the Past*, 6: 531-552.

Walker J.D. & Geissman J.W. (2009). GSA Geologic Time Scale. *GSA Today*, 19: 60-61.

Walker M., Head M.J., Berkelhammer M., Björck S., Cheng H., Cwynar L., Fisher D., Gkinis V., Long A., Lowe J., Newnham R., Rasmussen S.O. & Weiss H. (2018). Formal ratification of the subdivision of the Holocene Series/Epoch (Quaternary System/Period): two new Global Boundary Stratotype Sections and Points (GSSPs) and three new stages/subseries. *Episodes*, 41: 213-223.

Wallich G.C. (1877). Observations on the coccosphere. *Annals and Magazine of Natural History*, 19: 342-350.

Weber van Bosse A. (1901). Études sur les algues de l'Archipel Malaisien III. Note préliminaire sur les résultats algologiques de l'expédition du Siboga. *Annales du Jardin Botanique de Buitenzorg*, 17: 126-141.

Wright A.K. & Flower B.P. (2002). Surface and deep ocean circulation in subpolar North Atlantic during the mid-Pleistocene revolution. *Paleoceanography*, 17: 1068.
Zachariasse W.J., Zijderveld J.D.A., Langereis C.G., Hilgen F.J. & Verhallen P.J.J.M. (1989). Early late Pliocene biochronology and surface Water temperature variations in the Mediterranean. *Marine Micropaleontology*, 14: 339-355.

Zachariasse W.J., Gudjonsson L., Hilgen F.J., Langereis C.G., Lourens L.J., Verhallen P.J.J.M. & Zijderveld J.D.A. (1990). Late Gauss to Early Matuyama invasions of *Neogloboquadrina atlantica* in the Mediterranean and associated record of climatic change. *Paleoceanography*, 5: 239-252.

Zanola E., Bonomo S., Incarbona A., Di Stefano A., Distefano S., Ferretti P., Fornaciari E., Galeotti S., Macri P., Raffi I., Sabatino N., Speranza F., Sprovieri M., Di Stefano E., Sprovieri R., Rio D. & Capraro L. (2024). High-resolution climate variability across the Piacenzian/Gelasian boundary in the Monte San Nicola section (Sicily, Italy). *Quaternary Science Reviews*, 324: 108469.

Zijderveld J., Hilgen F., Langereis C., Verhallen P. & Zachariasse W. (1991). Integrated magnetostratigraphy and biostratigraphy of the upper Pliocene-lower Pleistocene from the Monte Singa and Crotone areas in Calabria, Italy. *Earth and Planetary Science Letters*, 107: 697-714.