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Pollen analyses have been carried out on the infilling deposits of Grotta Romanelli (Apulia, Italy), a reference site for the Middle and Upper Palaeolithic of Italy. The analysis focused on Terre rosse, a fine unit till now ascribed to an interstadial phase following the Würm acme, and on the uppermost unit (Terre brune), recently dated to the latest Late Pleistocene-Early Holocene. Despite the diffuse barrenness and low pollen concentration of many levels, pollen data from Grotta Romanelli gives insights into the palaeoenvironmental setting of the deposits and their chronological attribution. The presence of Olea in all levels of Terre rosse strongly suggests their attribution to the Last Interglacial (Eemian), during which this plant was diffused in the Mediterranean area. The Terre brune deposition occurred when the environment was open, with rare trees and shrubs and prevailing steppe elements. This association reflects the climatic conditions of the Lateglacial, with evidence of both the warm interstadial Bølling/Allerød and the cold stadial Younger Dryas. Mediterranean, mesophilous and riparian arboreal elements are present, especially in the Early Holocene levels. Comparison with modern pollen material allowed some fossil grains, found in high amounts and in clusters, to be tentatively ascribed to the species Crithmum maritimum (Apiaceae), Muscari comosum and to Asparagus maritimus/Ornithogalum (Asparagaceae). The significant occurrence of such entomophilous plants reveals differential transportation inside the cave and, since most of them are edible and/or have medicinal properties, suggests an intentional introduction and possible use during time, by both Neanderthals and modern humans.

The authors acknowledge the following funding sources: Max Planck Independent Research Group, Palaeo-Science and History Group (A.I., A.M. and C.V.); Estonian Research Council #PRG323, PUT1173 (A.Pos., T.R., N.S. and S.V.); European Research Council #FP7 263735 (A.Bro. and A.Plu.), #MSC 655659 (A.E.); Georgetown Environmental Initiative (T.N.); Latvian Council of Science #LZP-2020/2-0060 (N.S. and N.J.); LLNL-JRNL-820941 (I.T.); NSF award #GSS-1228126 (S.M.); Polish-Swiss Research Programme #013/2010 CLIMPEAT (M.Lam.), #086/2010 CLIMPOL (A.W.); Polish Ministry of Science and Higher Education #N N306 275635 (M.K.); Polish National Science Centre #2019/03/X/ST10/00849 (M.Lam.), #2015/17/B/ST10/01656 (M.Lam.), #2015/17/B/ST10/03430 (M.So.), #2018/31/B/ST10/02498 (M.So.), #N N304 319636 (A.W.); SCIEX #12.286 (K.Mar.); Spanish Ministry of Economy and Competitiveness #REDISCO-HAR2017-88035-P (J.A.L.S.); Spanish Ministry of Education, Culture and Sports #FPU16/00676 (R.L.L.); Swedish Research Council #421-2010-1570 (P.L.), #2018-01272 (F.C.L. and A.S.); Volkswagen Foundation Freigeist Fellowship Dantean Anomaly (M.B.), Spanish Ministry of Science and Innovation #RTI2018-101714-B-I00 (F.A.S. and D.A.S.), OP RDE, MEYS project #CZ.02.1.01/0.0/0.0/16_019/0000728 (P.P.)., The Black Death (1347–1352 ce) is the most renowned pandemic in human history, believed by many to have killed half of Europe’s population. However, despite advances in ancient DNA research that conclusively identified the pandemic’s causative agent (bacterium Yersinia pestis), our knowledge of the Black Death remains limited, based primarily on qualitative remarks in medieval written sources available for some areas of Western Europe. Here, we remedy this situation by applying a pioneering new approach, ‘big data palaeoecology’, which, starting from palynological data, evaluates the scale of the Black Death’s mortality on a regional scale across Europe. We collected pollen data on landscape change from 261 radiocarbon-dated coring sites (lakes and wetlands) located across 19 modern-day European countries. We used two independent methods of analysis to evaluate whether the changes we see in the landscape at the time of the Black Death agree with the hypothesis that a large portion of the population, upwards of half, died within a few years in the 21 historical regions we studied. While we can confirm that the Black Death had a devastating impact in some regions, we found that it had negligible or no impact in others. These inter-regional differences in the Black Death’s mortality across Europe demonstrate the significance of cultural, ecological, economic, societal and climatic factors that mediated the dissemination and impact of the disease. The complex interplay of these factors, along with the historical ecology of plague, should be a focus of future research on historical pandemics., Max Planck Independent Research Group, Palaeo-Science and History Group, Estonian Research Council PRG323 PUT1173, European Research Council (ERC) European Commission FP7 263735 MSC 655659, Georgetown Environmental Initiative, Latvian Ministry of Education and Science LZP-2020/2-0060 LLNL-JRNL-820941, National Science Foundation (NSF) GSS-1228126, Polish-Swiss Research Programme 013/2010 086/2010, Ministry of Science and Higher Education, Poland N306 275635, Polish National Science Centre 2019/03/X/ST10/00849 2015/17/B/ST10/01656 2015/17/B/ST10/03430 2018/31/B/ST10/02498 N N304 319636, SCIEX 12.286, Spanish Government REDISCO-HAR2017-88035-P FPU16/00676, Swedish Research Council, European Commission 421-2010-1570 2018-01272, Volkswagen Foundation Freigeist Fellowship Dantean Anomaly, Spanish Government RTI2018-101714-B-I00, OP RDE, MEYS project CZ.02.1.01/0.0/0.0/16_019/0000728

Mediterranean mid-altitude sites are critical for the survival of plant species allowing for elevational vegetation shifts in response to high-amplitude climate variability. Pollen records from the southern Balkans have underlined the importance of the region in preserving plant diversity over at least the last half a million years. So far, there are no records of vegetation and climate dynamics from Balkan refugia with an Early Pleistocene age. Here we present a unique palynological archive from such a refugium, the Lake Ohrid basin, recording continuously floristic diversity and vegetation succession under obliquity-paced climate oscillations. Palynological data are complemented by biomarker, diatom, carbonate isotope and sedimentological data to identify the mechanisms controlling shifts in the aquatic and terrestrial ecosystems within the lake and its catchment. The study interval encompasses four complete glacial-interglacial cycles (1365–1165 ka; MIS 43–35). Within the first 100 kyr of lake ontogeny, lake size and depth increase before the lake system enters a new equilibrium state as observed in a distinct shift in biotic communities and sediment composition. Several relict tree genera such as Cedrus, Tsuga, Carya, and Pterocarya played an important role in ecological succession cycles, while total relict abundance accounts for up to half of the total arboreal vegetation. The most prominent biome during interglacials is cool mixed evergreen needleleaf and deciduous broadleaf forests, while cool evergreen needleleaf forests dominate within glacials. A rather forested landscape with a remarkable plant diversity provide unique insights into Early Pleistocene ecosystem resilience and vegetation dynamics. © 2019

Archaeobotany is used to discover details on local land uses in prehistoric settlements developed during the middle and beginning of late Holocene. Six archaeological sites from four countries (Spain, Italy, Greece, and Turkey) have pollen and charcoal records showing clear signs of the agrarian systems that had developed in the Mediterranean basin during different cultural phases, from pre-Neolithic to Recent Bronze Age. A selected list of pollen taxa and sums, including cultivated trees, other woody species, crops and annual or perennial synanthropic plants are analysed for land use reconstructions. In general, cultivation has a lower image in palynology than forestry, and past land uses became visible when oakwoods were affected by human activities. On-site palynology allows us to recognise the first influence of humans even before it can be recognised in off-site sequences, and off-site sequences can allow us to determine the area of influence of a site. Neolithic and Bronze Age archaeological sites show similar land use dynamics implying oak exploitation, causing local deforestation, and cultivation of cereal fields in the area or around the site. Although a substantial difference makes the Neolithic influence quite distant from the Bronze Age impact, mixed systems of land exploitation emerged everywhere. Multiple land use activities exist (multifunctional landscapes) at the same time within the area of influence of a site. Since the Neolithic, people have adopted a diffuse pattern of land use involving a combination of diverse activities, using trees–crops–domesticated animals. The most recurrent combination included wood exploitation, field cultivation and animal breeding. The lesson from the past is that the multifunctional land use, combining sylvo-pastoral and crop farming mixed systems, has been widely adopted for millennia, being more sustainable than the monoculture and a promising way to develop our economy. © The Author(s) 2019.

Mediterranean climates are characterized by strong seasonal contrasts between dry summers and wet winters. Changes in winter rainfall are critical for regional socioeconomic development, but are difficult to simulate accurately1 and reconstruct on Quaternary timescales. This is partly because regional hydroclimate records that cover multiple glacial–interglacial cycles2,3 with different orbital geometries, global ice volume and atmospheric greenhouse gas concentrations are scarce. Moreover, the underlying mechanisms of change and their persistence remain unexplored. Here we show that, over the past 1.36 million years, wet winters in the northcentral Mediterranean tend to occur with high contrasts in local, seasonal insolation and a vigorous African summer monsoon. Our proxy time series from Lake Ohrid on the Balkan Peninsula, together with a 784,000-year transient climate model hindcast, suggest that increased sea surface temperatures amplify local cyclone development and refuel North Atlantic low-pressure systems that enter the Mediterranean during phases of low continental ice volume and high concentrations of atmospheric greenhouse gases. A comparison with modern reanalysis data shows that current drivers of the amount of rainfall in the Mediterranean share some similarities to those that drive the reconstructed increases in precipitation. Our data cover multiple insolation maxima and are therefore an important benchmark for testing climate model performance. © 2019, The Author(s), under exclusive licence to Springer Nature Limited.

International audience

The 7.5 m long SK13 sediment core, drilled at Lake Shkodra (Albania) bottom depth of 7 m in the central southern part of the lake, was selected for multidisciplinary analysis. Ostracods, Characeae, pollens, and stable isotopes were studied with the aim to reconstruct the past biodiversity and the palaeoenvironmental and palaeoclimatic changes occurred during the Middle-Late Holocene. The chronological framework of SK13 was already known and well constrained through radiometric datings. The sediment core age spans from 4560 cal year BP to present. Thirteen ostracod and five charophyte species were identified. Among them, the ostracods Candona montenigrina and Limnocythere scutariense are endemic of the lake; Candona meridionalis and Paralimnocythere georgevitschi have been collected for the first time in the Lake Shkodra while formerly were considered endemic respectively of Lake Dojran and Lake Prespa; Metacypris cordata, ?Carpathocandona sp., and Cyclocypris sp. were never recorded in the lake and were found only in the lower portion of the sediment core. Among the charopytes, Lychnothamnus barbatus and N. hyalina are recorded for the first time in the lake and occurs with high frequency throughout interval A of core. The faunal composition is quite homogeneous, with the percentages of the different species varying along the sediment core. The main change occurs between 1274 and 1197 cal yr BP, where charophytes disappear abruptly as well as 8 ostracod species out of 13, and the frequency of the remaining 5 species dramatically increases. Moreover at that moment, it occurs also the appearance of a peculiar not-tuberculate morphotypes of Ilyocypris monstrifica. The micropaleontological data suggest a decrease of the lake biodiversity since around 1200 cal years BP linked to the transition between an ancient marshland to a lacustrine environment. This drastic change seems to be independent from any global or local climate changes and could be related to the co-occurrence of two phenomena: 1) the Adriatic sea level rise during the Middle and Late Holocene, that that would have uplifted the base level of the River Bojana causing a decrease of its discharge and, consequently, reducing the drainage of the water from the Shkodra swamp; 2) the periodical change of the River Drin course over the last five centuries, which, when deviated into the Bojana river, blocked the River Bojana outflow.

Climate evolution of the Mediterranean region during the Holocene exhibits strong spatial and temporal variability, which is notoriously difficult for models to reproduce. We propose here a new proxy-based climate synthesis synthesis and its comparison - at a regional ( ~ 100 km) level - with a regional climate model to examine (i) opposing northern and southern precipitation regimes and (ii) an east-to-west precipitation dipole during the Holocene across the Mediterranean basin. Using precipitation estimates inferred from marine and terrestrial pollen archives, we focus on the early to mid-Holocene (8000 to 6000 cal yr BP) and the late Holocene (4000 to 2000 cal yr BP), to test these hypotheses on a Mediterranean-wide scale. Special attention was given to the reconstruction of season-specific climate information, notably summer and winter precipitation. The reconstructed climatic trends corroborate the north-south partition of precipitation regimes during the Holocene. During the early Holocene, relatively wet conditions occurred in the south-central and eastern Mediterranean regions, while drier conditions prevailed from 45° N northwards. These patterns then reverse during the late Holocene. With regard to the existence of a west-east precipitation dipole during the Holocene, our results show that the strength of this dipole is strongly linked to the reconstructed seasonal parameter; early-Holocene summers show a clear east-west division, with summer precipitation having been highest in Greece and the eastern Mediterranean and lowest over Italy and the western Mediterranean. Summer precipitation in the east remained above modern values, even during the late-Holocene interval. In contrast, winter precipitation signals are less spatially coherent during the early Holocene but low precipitation is evidenced during the late Holocene. A general drying trend occurred from the early to late Holocene, particularly in the central and eastern Mediterranean. For the same time intervals, pollen-inferred precipitation estimates were compared with model outputs, based on a regional-scale downscaling (HadRM3) of a set of global climate-model simulations (HadAM3). The highresolution detail achieved through the downscaling is intended to enable a better comparison between site-based paleo-reconstructions and gridded model data in the complex terrain of the Mediterranean; the model outputs and polleninferred precipitation estimates show some overall correspondence, though modeled changes are small and at the absolute margins of statistical significance. There are suggestions that the eastern Mediterranean experienced wetter summer conditions than present during the early and late Holocene; the drying trend in winter from the early to the late Holocene also appears to be simulated. The use of this highresolution regional climate model highlights how the inherently patchy nature of climate signals and paleo-records in the Mediterranean basin may lead to local signals that are much stronger than the large-scale pattern would suggest. Nevertheless, the east-to-west division in summer precipitation seems more marked in the pollen reconstruction than in the model outputs. The footprint of the anomalies (like today, or dry winters and wet summers) has some similarities to modern analogue atmospheric circulation patterns associated with a strong westerly circulation in winter (positive Arctic Oscillation-North Atlantic Oscillation (AO-NAO)) and a weak westerly circulation in summer associated with anticyclonic blocking; however, there also remain important differences between the paleo-simulations and these analogues. The regional climate model, consistent with other global models, does not suggest an extension of the African summer monsoon into the Mediterranean. Therefore, the extent to which summer monsoonal precipitation may have existed in the southern and eastern Mediterranean during the mid- Holocene remains an outstanding question. © Author(s) 2017.

This study reviews and synthesises existing information generated within the SCOPSCO (Scientific Collaboration on Past Speciation Conditions in Lake Ohrid) deep drilling project. The four main aims of the project are to infer (i) the age and origin of Lake Ohrid (Former Yugoslav Republic of Macedonia/Republic of Albania), (ii) its regional seismotectonic history, (iii) volcanic activity and climate change in the central northern Mediterranean region, and (iv) the influence of major geological events on the evolution of its endemic species. The Ohrid basin formed by transtension during the Miocene, opened during the Pliocene and Pleistocene, and the lake established de novo in the still relatively narrow valley between 1.9 and 1.3 Ma. The lake history is recorded in a 584 m long sediment sequence, which was recovered within the framework of the International Continental Scientific Drilling Program (ICDP) from the central part (DEEP site) of the lake in spring 2013. To date, 54 tephra and cryptotephra horizons have been found in the upper 460 m of this sequence. Tephrochronology and tuning biogeochemical proxy data to orbital parameters revealed that the upper 247.8 m represent the last 637 kyr. The multi-proxy data set covering these 637 kyr indicates long-term variability. Some proxies show a change from generally cooler and wetter to drier and warmer glacial and interglacial periods around 300 ka. Short-term environmental change caused, for example, by tephra deposition or the climatic impact of millennial-scale Dansgaard-Oeschger and Heinrich events are superimposed on the long-term trends. Evolutionary studies on the extant fauna indicate that Lake Ohrid was not a refugial area for regional freshwater animals. This differs from the surrounding catchment, where the mountainous setting with relatively high water availability provided a refuge for temperate and montane trees during the relatively cold and dry glacial periods. Although Lake Ohrid experienced significant environmental change over the last 637 kyr, preliminary molecular data from extant microgastropod species do not indicate significant changes in diversification rate during this period. The reasons for this constant rate remain largely unknown, but a possible lack of environmentally induced extinction events in Lake Ohrid and/or the high resilience of the ecosystems may have played a role. © Author(s) 2017.

International audience; Lake Ohrid (Balkan peninsula) is the oldest European extant lake and one of the deepest and largest. Such a unique, terrestrial natural archive is especially relevant for both paleoenvironmental and paleoclimatic reconstructions but also for genetic studies. In the frame of the International Continental Scientific Drilling Program (ICDP), a deep drilling campaign was carried out within the scope of the Scientific Collaboration on Past Speciation Conditions in Lake Ohrid (SCOPSCO) project in 2013. Here, we present the summary of palynological analyses carried out in the upper 200 m of the overall 569 m long DEEP site sediment succession from the central part of the lake. These studies, performed by an international palynological team, document the main floristic, vegetation and climate changes during the last ca 500 kyr, at a millennial-scale resolution (~1.6 kyr). The continuous sediment infill permitted to trace multiple non-forested/ forested phases as a response to Glacial/Interglacial cycles as well as to sub-Milankovitch climate changes. The pollen record, corresponding with marine isotope stages MIS 13 to MIS 1, points to a progressive change from cooler and wetter to warmer and drier interglacials. New palynological studies are underway to reconstruct vegetational and climatic conditions over older intervals as well as to obtain high resolution data for some key intervals such as MIS 5-6, MIS 11-12, MIS 35-42. The complete record of changes in flora composition and vegetation during both glacials and interglacials will furnish indispensable insights for understanding the role of refugia, ecosystem resilience and maintenance of terrestrial biodiversity in the Mediterranean area.

Lake Ohrid is located at the border between FYROM (Former Yugoslavian Republic of Macedonia) and Albania and formed during the latest phases of Alpine orogenesis. It is the deepest, the largest and the oldest tectonic lake in Europe. To better understand the paleoclimatic and paleoenvironmental evolution of Lake Ohrid, deep drilling was carried out in 2013 within the framework of the Scientific Collaboration on Past Speciation Conditions (SCOPSCO) project that was funded by the International Continental Scientific Drilling Program (ICDP). Preliminary results indicate that lacustrine sedimentation of Lake Ohrid started between 1.2 and 1.9 Ma ago. Here we present new pollen data (selected percentage and concentration taxa/groups) of the uppermost ∼ 200 m of the 569 m long DEEP core drilled in the depocentre of Lake Ohrid. The study is the fruit of a cooperative work carried out in several European palynological laboratories. The age model of this part of the core is based on 10 tephra layers and on tuning of biogeochemical proxy data to orbital parameters. According to the age model, the studied sequence covers the last ∼ 500 000 years at a millennial-scale resolution ( ∼ 1.6 ka) and records the major vegetation and climate changes that occurred during the last 12 (13 only pro parte) marine isotope stages (MIS). Our results indicate that there is a general good correspondence between forested/non-forested periods and glacial-interglacial cycles of the marine isotope stratigraphy. The record shows a progressive change from cooler and wetter to warmer and drier interglacial conditions. This shift in temperature and moisture availability is visible also in vegetation during glacial periods. The period corresponding to MIS11 (pollen assemblage zone OD-10, 428-368 ka BP) is dominated by montane trees such as conifers. Mesophilous elements such as deciduous and semi-deciduous oaks dominate forest periods of MIS5 (PASZ OD-3, 129-70 ka BP) and MIS1 (PASZ OD-1, 14 ka BP to present). Moreover, MIS7 (PASZ OD-6, 245-190 ka) shows a very high interglacial variability, with alternating expansions of montane and mesophilous arboreal taxa. Grasslands (open vegetation formations requiring relatively humid conditions) characterize the earlier glacial phases of MIS12 (PASZ OD-12, 488-459 ka), MIS10 (corresponding to the central part of PASZ OD-10, 428-366 ka) and MIS8 (PASZ OD-7, 288-245 ka). Steppes (open vegetation formations typical of dry environments) prevail during MIS6 (OD-5 and OD-4, 190-129 ka) and during MIS4-2 (PASZ OD-2, 70-14 ka). Our palynological results support the notion that Lake Ohrid has been a refugium area for both temperate and montane trees during glacials. Closer comparisons with other long southern European and Near Eastern pollen records will be achieved through ongoing high-resolution studies. © 2016 Author(s).

Lake Ohrid (Balkan peninsula) is the oldest European extant lake and one of the deepest and largest. Such a unique, terrestrial natural archive is especially relevant for both paleoenvironmental and paleoclimatic reconstructions but also for genetic studies. In the frame of the International Continental Scientific Drilling Program (ICDP), a deep drilling campaign was carried out within the scope of the Scientific Collaboration on Past Speciation Conditions in Lake Ohrid (SCOPSCO) project in 2013. Here, we present the summary of palynological analyses carried out in the upper 200 m of the overall 569 m long DEEP site sediment succession from the central part of the lake. These studies, performed by an international palynological team, document the main floristic, vegetation and climate changes during the last ca 500 kyr, at a millennial-scale resolution (~1.6 kyr). The continuous sediment infill permitted to trace multiple non-forested/ forested phases as a response to Glacial/Interglacial cycles as well as to sub-Milankovitch climate changes. The pollen record, corresponding with marine isotope stages MIS 13 to MIS 1, points to a progressive change from cooler and wetter to warmer and drier interglacials. New palynological studies are underway to reconstruct vegetational and climatic conditions over older intervals as well as to obtain high resolution data for some key intervals such as MIS 5-6, MIS 11-12, MIS 35-42. The complete record of changes in flora composition and vegetation during both glacials and interglacials will furnish indispensable insights for understanding the role of refugia, ecosystem resilience and maintenance of terrestrial biodiversity in the Mediterranean area.

This paper reviews the methodological and practical issues relevant to the ways in which natural scientists, historians and archaeologists may collaborate in the study of past climatic changes in the Mediterranean basin. We begin by discussing the methodologies of these three disciplines in the context of the consilience debate, that is, attempts to unify different research methodologies that address similar problems. We demonstrate that there are a number of similarities in the fundamental methodology between history, archaeology, and the natural sciences that deal with the past ("palaeoenvironmental sciences"), due to their common interest in studying societal and environmental phenomena that no longer exist. The three research traditions, for instance, employ specific narrative structures as a means of communicating research results. We thus present and compare the narratives characteristic of each discipline; in order to engage in fruitful interdisciplinary exchange, we must first understand how each deals with the societal impacts of climatic change. In the second part of the paper, we focus our discussion on the four major practical issues that hinder communication between the three disciplines. These include terminological misunderstandings, problems relevant to project design, divergences in publication cultures, and differing views on the impact of research. Among other recommendations, we suggest that scholars from the three disciplines should aim to create a joint publication culture, which should also appeal to a wider public, both inside and outside of academia. © 2015 The Authors.

During September 2003 several cores were drilled from the Albanian side of Lake Shkodra. Among them, a 7,8 m long composite core (SK13), drilled at a lake bottom depth of 7 m in the central southern part of the lake (42°6’11.65”N, 19°25’18.96”E) has been selected for multidisciplinary analysis. Ostracods, characeae, pollen, CaCO3 content and stable isotopes were studied with the aim to reconstruct the palaeoenvironmental and palaeoclimatic changes occurred during the Late Holocene. The chronological framework of SK13 was established through the recognition of four well-dated tephra layers and four 14C accelerator mass spectrometry measurements; the sedimentation rate has been calculated as spanning between 1.1 and 3.0 mm/yr in the different portions of the composite sediment core (SULPIZIO et al., 2010, VAN WELDEN et al., 2008).

A new global synthesis and biomization of long (>40 kyr) pollen-data records is presented, and used with simulations from the HadCM3 and FAMOUS climate models to analyse the dynamics of the global terrestrial biosphere and carbon storage over the last glacial–interglacial cycle. Global modelled (BIOME4) biome distributions over time generally agree well with those inferred from pollen data. The two climate models show good agreement in global net primary productivity (NPP). NPP is strongly influenced by atmospheric carbon dioxide (CO2) concentrations through CO2 fertilization. The combined effects of modelled changes in vegetation and (via a simple model) soil carbon result in a global terrestrial carbon storage at the Last Glacial Maximum that is 210–470 Pg C less than in pre-industrial time. Without the contribution from exposed glacial continental shelves the reduction would be larger, 330–960 Pg C. Other intervals of low terrestrial carbon storage include stadial intervals at 108 and 85 ka BP, and between 60 and 65 ka BP during Marine Isotope Stage 4. Terrestrial carbon storage, determined by the balance of global NPP and decomposition, influences the stable carbon isotope composition (δ13C) of seawater because terrestrial organic carbon is depleted in 13C. Using a simple carbon-isotope mass balance equation we find agreement in trends between modelled ocean δ13C based on modelled land carbon storage, and palaeo-archives of ocean δ13C, confirming that terrestrial carbon storage variations may be important drivers of ocean δ13C changes.