Eemian and Early Weichselian vegetation and climate history in Central Europe: A case study from the Klinge section (Lusatia, eastern Germany)

The paper presents vegetation and climate reconstructions for Eemian and Early Weichselian in Central Europe based on detailed pollen data of the Klinge section. The sediments exposed in this profile were accumulated from the Saalian termination up to the Early Weichselian glaciation. A succession of forest communities, typical for the Eemian interglacial has been recorded. Pollen data indicate some oscillations during the very end of the Eemian (biozone E7). The climatic reconstructions show the heat supply rising gradually from the beginning of the interglacial to its middle part. A long-term cooling trend in January temperature was reconstructed for the second half of the interglacial. The difference between the optimum (zone E4b) and Pinus-Picea-Abies zone (E6) amounts to 4 °C, during the Pinus zone (E7) the temperature further decreased by 8 °C. The changes in July temperature were not so dramatic (about 3 °C).     

A Late-Pleistocene pollen sequence from the Jammertal, south-western Germany with particular reference to location and altitude as factors determining Eemian forest composition

Pollen analytical data from a 23-m core taken in the Jammertal subglacial basin, near Biberach an der Riss, has given a continuous sequence from the Rissian late-glacial to the beginning of the Mid-Würmian. The Eemian interglacial is completely preserved and with a high temporal resolution. Two interstadials, Stafflangen I and II, are recognised in the Lower Würmian. These two woodlands were dominant byPinus andPicea and there was a small but significant (up to 12%) component of thermophilous deciduous tree pollen. Stafflangen I and II are correlated with St. Germain I and II as known from Grande Pile. The north-eastern areal boundary of thermophilous deciduous trees lies between Jammertal and Grande Pile. Furthermore, it was possible for the first time to define and describe altitudinal zonation in Eemian vegetation. During the Eemian, the altitudinal boundary of shade-tolerant deciduous trees (Carpinus, Abies andPicea) lay at ca. 625 m in Upper Swabia and the Swiss Alpine foreland, and at ca. 560 m in Upper Bavaria to the east. The sites, which were dominated byCarpinus occur below this limit whereasAbies orPicea show a preference for habitats at higher altitudes.     

A 200-ka pollen and oxygen-isotopic record from two sediment cores from the eastern Arabian Sea

We present here a pollen record from the northern Indian Ocean that goes back to 200 ka, the boundary between marine isotopic stages (MIS) 7 and 6. Pollen, oxygen-isotopic composition and organic carbon have been examined for two sediment cores from the eastern Arabian Sea (15°02′N and 71°41′E, 13°16′N and 71°00′E), to reconstruct the long-term palaeoclimate and palaeovegetation of the Indian subcontinent. Oxygen-isotope data suggest that glacial periods (MIS 2, 4 and 6) are characterised by low precipitation because of a weak southwest monsoon and a strong northeast monsoon. In contrast, interglacial periods (MIS 1, 3 and 5) are marked by high fresh water input resulting from a strong southwest monsoon. During the last glacial-interglacial cycle, sea-surface temperature and surface salinity changed as a result of variations in the evaporation-precipitation (E-P) balance. Throughout the core, the dominant pollen types are Poaceae and Chenopodiaceae and/or Amaranthaceae (>50%). Their dominance during glacial periods (MIS 2, 4 and 6) suggests that the climate was cold and dry. This dominance is also suggestive of salinity-tolerant vegetation colonizing large areas near seashore due to lower sea level or high E-P conditions. During interglacial periods (MIS 1, 3 and 5) when there was high precipitation, the arid taxa were sparse, whereas Poaceae and Piperaceae were abundant. Hence, Chenopodiaceae and/or Amaranthaceae and Artemisia are suggestive of cold and arid/semi-arid climate, and Poaceae and Piperaceae of warm and wet conditions. Mangrove pollen is not well represented in the cores.     

Vegetation, climate and palaeoaltitude reconstructions of the Eastern Alps during the Miocene based on pollen records from Austria, Central Europe

Aim To reconstruct the flora, vegetation, climate and palaeoaltitude during the Miocene (23.03–5.33 Ma) in Central Europe. Location Six outcrop sections located in different basins of the Central Paratethys in Austria. Methods Pollen analysis was used for the reconstruction of the vegetation and climate. The altitude of the Eastern Alps that are adjacent to the Alpine Foreland and Vienna basins has been estimated using a new quantification method based on pollen data. This method uses biogeographical and climatological criteria such as the composition of the modern vegetation belts in the European mountains and Miocene annual temperature estimates obtained from fossil pollen data. Results Pollen changes from Early to Late Miocene have been observed. The vegetation during the Burdigalian and Langhian (20.43–13.65 Ma) was dominated by thermophilous elements such as evergreen trees, typical of a present‐day evergreen rain forest at low altitudes (i.e. south‐eastern China). During the Serravallian and Tortonian (13.65–7.25 Ma) several thermophilous elements strongly decreased, and some disappeared from the Central European region. This kind of vegetation was progressively substituted by one enriched in deciduous and mesothermic plants. Middle‐altitude (Cathaya, Cedrus and Tsuga) and high‐altitude (Abies and Picea) conifers increased considerably during the Langhian and later on during the Serravallian and Tortonian. Main conclusions Pollen changes are related to climatic changes and to the uplift of the Alpine massifs. The vegetation during the Burdigalian and Langhian reflects the Miocene climatic optimum. The decrease in thermophilous plants during the Serravallian and Tortonian can be interpreted as a climatic cooling and can be correlated with global and regional climatic changes. This study shows that the palaeoaltitude of the eastern part of the Eastern Alps during the Burdigalian was not high enough for Abies and Picea to form a forest. Therefore, we inferred that the summits of most of the mountains would have been less than 1800 m. The substantial increase of middle‐ and high‐altitude conifers in the pollen spectra suggests that the uplift rate increased during the Langhian in this region. Based on higher palaeoaltitude estimations for the pollen floras from the studied sections of Austria, we infer that the uplift of the easternmost part of the Alpine chain continued during the Serravallian and Tortonian.     

Late Miocene vegetation and climate reconstruction based on pollen data from the Sofia Basin (West Bulgaria)

The analysis of fossil palynomorh assemblages in the Late Miocene freshwater sediments of the Sofia Basin (West Bulgaria) was done to collect data on the vegetation and climate dynamics during the Late Miocene. On the basis of pollen data, we described the main palaeocomunities developed in the region. The mixed mesophytic forests dominated the vegetation in which species of Quercus, Ulmus, Zelkova, Fagus, Carpinus, Betula, Castanea, Corylus, Pterocarya, Carya, Juglans, and Eucommia played important roles. Swamp forests were also recorded, including Taxodiaceae, Alnus, Glyptostrobus, Nyssa, and Myrica. Herbaceous vegetation was distributed in the middle part of the section, with a maximum of 35.5%. The vegetation dynamic passes through several phases, which were associated with changes in paleoclimate and palaeoecological conditions. Coexistence Approach (CA) was applied to palynological data to calculate four climatic parameters. The values of coexistence intervals for mean annual temperatures are 13.6–16.6 °C, with winter temperatures being 3.7–6.6 °C and summer temperatures being 23.6–27.8 °C. Mean annual precipitation ranged most frequently between 828 and 1308 mm. The palaeoclimatic reconstruction illustrates existence of a warm-temperate and relatively humid climate with higher mean annual temperature than the present day climate.     

A diseased Panthera leo spelaea (Goldfuss 1810) lioness from a forest elephant graveyard in the Late Pleistocene (Eemian) interglacial lake at Neumark-Nord,central Germany

A solitary articulated skeleton of a middle-aged and diseased Panthera leo spelaea lioness from the Eemian interglacial has been found amongst numerous articulated skeletons of Palaeoloxodon antiquus forest elephants, in sediments from a small, shallow lake at Neumark-Nord in central Germany, which has Neanderthal settlements along its shoreline. Several pathologies such as a fibula fracture, arthritis in one of the front legs and a lost canine tooth with associated maxillary inflammation and dissolution made the lioness vulnerable to other predators such as hyenas, whose presence is indicated by their bones, coprolites and many scavenging marks on the elephant skeletons and on a femur from a male lion. The scavenging of hyenas and lions at this site is commonly documented by canine bite marks on the joints of elephant bones. Bite and scratch marks on the ventral vertebral columns and pelvises of two P. antiquus forest elephant skeletons suggest that the intestines and inner organs may have been consumed by large predators, as is commonly the case with modern African lions feeding on elephants. The weak and diseased lioness may possibly have been killed during antagonistic battles between hyenas and lions over their larger prey.     

A new contribution to the Holocene vegetation history of the West African Sahel: pollen from Oursi,Burkina Faso and charcoal from three sites in northeast Nigeria

A pollen diagram from Oursi in Burkina Faso is compared with anthracological (charcoal analysis) results from three sites in northeast Nigeria (Konduga, Gajiganna, Lantewa). The present-day vegetation at all four sites is Sahelian or Sahelo-Sudanian and under heavy human impact. At Oursi, a closed grassland with only few trees and almost no Sudanian elements can be reconstructed for the middle Holocene. At the Nigerian sites, on the other hand, Sudanian woody plants were present during this period. We assume that the Sahel was not a uniform zone during the middle Holocene but rather a mosaic of different vegetation types according to local site conditions. In the light of these results, a simple model of latitudinally shifting vegetation zones is not applicable. Around 3000 B.P. the closed grassland at Oursi was opened by agro-pastoral activities, and at Gajiganna, plants characteristic of pasture lands can be directly related with the presence of cattle. Human impact seems to have been the dominant factor in the vegetation history of the Sahel from 3000 B.P. until today, masking possible effects of climatic change.     

Towards the reconstruction of the Holocene vegetation history of Lower Provence: two new pollen profiles from Marais des Baux

Two new¹⁴C-dated Holocene pollen profiles from Marais des Baux, Bouches-du-Rhône, France, are presented. The record begins in the Younger Dryas, when the vegetation consisted mainly of grasses and mugwort (Artemisia). The Preboreal was marked by a transitory expansion of pine forests and was followed by the establishment of a rich deciduous oak-forest that included hazel (Corylus) and elm (Ulmus). During the Boreal, hazel played a dominant role within the oak-forest. The oak forests, which includedQuercus ilex, achieved a major expansion during the Atlantic period. The Subboreal was characterised by the regional establishment of, firstly, fir (Abies) and then beech (Fagus). The spread and expansion of beech coincides with the first clear evidence for farming. Agricultural activities brought about the decline of deciduous oak-forest. During the Subatlantic, forests in the vicinity of Marais des Baux were cleared for farming. Cereal growing, which included rye cultivation, was of considerable importance. Three noteworthy characteristics that serve to differentiate the Holocene vegetation history of the low-lying Provence region from other French regions are as follows: 1) the early establishment (from the onset of Preboreal) of low altitude mixed forest; 2) the expansion during the Subboreal of fir and beech in low altitude areas with a Mediterranean climate and, 3) the exceptional taxonomic richness of the pollen assemblages (120 identified taxa) and the presence of borealalpine and Euro-Siberian taxa that no longer exist in Lower Provence.     

Reconstruction of the vegetation at A.D. 915 at Ohse-yachi Mire, northern Japan, from pollen, present-day vegetation and tephra data

The Towada-a (To-a) Tephra from A.D. 915 allowed reconstruction of the local and regional vegetation at and near Ohse-yachi Mire from pollen samples taken just below the tephra. In earlier studies the tephra was dated from ¹⁴C data, stratigraphy at archaeological sites and a historical record. Twelve cores along two perpendicular transects yielded pollen from modern and pre-tephra samples. The modern pollen data were compared with the modern vegetation, and used to interpret the pollen data from A.D. 915. Ericaceae, Drosera, Sanguisorba, other Rosaceae, and Sphagnum were concluded to be local elements. Ohse-yachi Mire was a wet Sphagnum bog in A.D. 915, with Sphagnum dominating the central part of the mire. The present stand of scrub was not yet established, but there was a stand of Pinus pumila scrub. The bog subsequently became drier and changed to the present-day mixed Moliniopsis-Sphagnum bog.     

Miocene to Pliocene vegetation reconstruction and climate estimates in the Iberian Peninsula from pollen data

Pollen analysis of Miocene and Pliocene sediments from the Iberian Peninsula shows a progressive reduction in plant diversity through time caused by the disappearance of thermophilous and high-water requirement plants. In addition, an increase in warm-temperate (mesothermic), seasonal-adapted “Mediterranean” taxa, high-elevation conifers and herbs (mainly Artemisia) occurred during the Middle and Late Miocene and Pliocene. This has mainly been interpreted as a response of the vegetation to global and regional processes, including climate cooling related to the development of the East Antarctic Ice Sheet and then the onset of the Arctic Ice Sheet, uplift of regional mountains related to the Alpine uplift and the progressive movement of Eurasia towards northern latitudes as a result of the northwards subduction of Africa. The development of steppe-like vegetation in southern Iberia is ancient and probably started during the Oligocene. The onset of a contrasted seasonality in temperature during the Mid-Pliocene superimposed on the pre-existing seasonality in precipitation, the annual length of which increased southward. The Mediterranean climatic rhythm (summer drought) began about 3.4 Ma and caused the individualization of modern Mediterranean ecosystems. Quaternary-type Mediterranean climatic fluctuations started at 2.6 Ma (Gelasian) resulting in repeated steppe vs. forest alternations. A latitudinal climatic gradient between the southern and the northern parts of the Iberian Peninsula existed since the Middle Miocene.     

Six years of observation of airborne and deposited pollen in central European Russia: first results

The aim of the study was to compare the pollen spectra recorded in rural and urban environments in order to examine whether pollen data obtained in the urban environment could also represent the situation in surrounding rural areas and how urban planting affects the regional pollen spectrum. Daily airborne pollen observations were performed with a volumetric Hirst type trap in Moscow (Russia). Modified Tauber traps were located in the Moscow region. The Tauber traps were installed and treated according to Pollen Monitoring Programme (PMP) guidelines. Spearman correlation coefficients were determined between the percentage of total pollen recorded in the Tauber and Hirst type samplers. The data showed that the Tauber spectra are characterised by higher taxonomic variability, mainly due to the greater occurrence of local herbaceous plants. Pollen types registered by Hirst type samplers were predominantly represented by anemophilous arboreal species, used in urban planting. A significant correlation for annual sums was only observed for Betula pollen, the most abundant component in both sampler types. Data obtained in big cities for dominant taxa can be extrapolated to the surrounding rural territories. The simultaneous use of volumetric and gravimetric samplers can be very useful for palaeo-ecological studies to evaluate the presence of local and transported pollen.     

Taphonomy in the light of intrinsic shell properties and life habits: Marine bivalves from the Eemian of northern Russia

A taphonomic analysis of the bivalvesArctica islandica (Arcticidae),Astarte borealis (Astartidae),Mytilus edulis (Mytilidae), andSpisula elliptica (Mactridae) from shallow-marine last interglacial Sediments exposed along the Pyoza river, Arkhangelsk region, shows that they differ in preservation, probably because of differences in shell shape, shell structures, and life habits. The Shells indicate that the temporal sequence of taphonomic processes was as follows: (1) pre-mortem bioerosion and dissolution; (2) post-mortem bioerosion; (3) abrasion, disarticulation, and fragmentation; and (4) dissolution. A new graphic Illustration, the taphonomic constituent diagram (TCD), is proposed to illustrate the sequences of taphonomic processes. It is inspired by the ichnofabric constituent diagram used in ichnology and integrates the shell surface coverage of different taphonomic features by graphically plotting them against relative time. The taphonomic constituent diagram may display differences in the Chronologie order of paleoenvironmental processes. Also, the diagram enables a combination of both microscopic and macroscopic taphonomic features, and eases comparative studies of fossil assemblages.      

Mid to late Holocene vegetation and land use history in the Weald of south-eastern England: multiple pollen profiles from the Rye area

The High Weald is an unusually well-wooded area in southern England. A high proportion of this woodland is ancient, being formerly exploited as seasonal pasture and coppice. Multiple pollen profiles from the Rye area have been used to elucidate the origins of this cultural landscape. By combining sites with small and large pollen source areas, both local and regional patterns of vegetation change have been determined. The mid-Holocene Tilia-dominated woodlands were subjected to temporary clearance as early as the Neolithic. This woodland was more extensively exploited over a ca. 700 year period from the beginning of the Bronze Age. The main elements of the modern landscape (woodland, pasture and limited cultivation) can be traced back to a more intensive phase of human activity, which commenced in the late Bronze Age. A regional increase in Fagus sylvatica pollen ca. 750 B.C. probably reflects the use of the Wealden woods for pasturage. There is no palynological evidence that the fuel demands of the Roman iron industry resulted in widespread woodland destruction. The early Anglo-Saxon period appears to have been one of land-use continuity, with a second increase in Fagus pollen at ca. A.D. 700 corresponding to historical evidence for the presence of wood-pastures in the Weald.     

A contribution to the ultrastructural knowledge of the pollen exine in subtribe Inulinae (Inuleae, Asteraceae)

To better understand the relationships within the Asteroideae and Inuleae, the structure of the pollen exine was investigated in seven genera and nine species of the subtribe Inulinae using LM, TEM and SEM. All taxa have a senecioid pattern of exine. The tectal complex consists of three main layers that differ in thickness and morphology: a tectum, a columellar layer, and a layer consisting of the basal region of the columellae. The absence or the vestigial condition of the foramina is considered as a plesiomorphy within the Asteroideae. All taxa have a complex apertural system that consists of an ecto-, a meso-, and an endoaperture. These apertures intersect respectively the tectal complex, the foot layer and the upper part of the endexine, and the inner layer of the endexine. A continuous transition among the different species of Inulinae was found for all the quantitative characters examined. This relative homogeneity of the pollen morphological characters enhances the naturality of the subtribe Inulinae.     

Modern pollen/land-use relationships as an aid in the reconstruction of past land-uses and cultural landscapes: an example from south Sweden

An on-going, integrated project concerning modern pollen/vegetation/land-use relationships in south Sweden is outlined. Some preliminary results and their potential uses in palaeoecological reconstructions are discussed. Moss polsters were collected from 92 examples of vegetation/land-use areas of south Sweden (e.g. nonfertilized grazed areas, burned and grazed heaths, traditionally managed fodder-producing meadows, etc.). A total of 23 land-use (e.g. management type) and environment variables (e.g. soil chemistry) is available for 84 sites. The 84 sample data-set was analysed by canonical correspondence analysis and associated statistical testing using Monte Carlo permutation tests to explore and test patterns of modern pollen variation in relation to the environment, and by weighted averaging regression and calibration to derive local-scale environmental and land-use reconstructions from fossil pollen assemblages preserved in a soil profile.     

A 2700-year high resolution pollen record of climate change from varved Sugan Lake in the Qaidam Basin, northeastern Tibetan Plateau

A 2700-year high resolution pollen record from annually-varved Sugan Lake in the Qaidam Basin at 2793 m a.s.l was obtained to examine vegetation and climatic change on the NE Tibetan Plateau. Pollen data shows that Sugan Basin was constantly covered by open desert-steppe vegetation dominated by Chenopodiaceae, Artemisia, Poaceae and Ephedra. However, large variations in Artemisia/Chenopodiaceae (A/C) ratios suggest regional moisture fluctuations over the last 2700 years, including a dry and relatively stable climate prior to 300 AD, relatively wet climate from 300 to 1200 AD with variability during 1100-1200 AD, and unstable climate since 1200 AD with relatively moister climate during 1250-1400 AD and 1700-1800 AD. However, other proxies (varve thickness, Chironomid taxa, isotopes of oxygen in precipitated carbonate) show fresher water when regional moisture was lower inferred from A/C ratio. This inconsistency suggests the possible difference of in-lake lithology/environment and regional moisture change. Fresh water into the lake from ice melting on the surrounding mountains might have contributed to the in-lake lithology and environment variation. The effective moisture changes in the Basin are in opposite phases to snow accumulation records from Dunde ice core (5325 m a.s.l) and to the monsoon intensity inferred from Dongge Cave, suggesting that the regional topography might have played an important role in mediating moisture changes at regional scale. Pollen data from Sugan Lake shows the shift of moisture at 1200 AD, from stable to variable conditions. This event is well correlated with other paleoclimate proxies in China and other parts of the world; however, the mechanisms behind these patterns require further investigation.     

The Massaciuccoli Holocene pollen sequence and the vegetation history of the coastal plains by the Mar Ligure (Tuscany and Liguria,Italy)

A 90 m deep core from the Massaciuccoli lake basin (Tuscany, Italy) offers new material for improving the knowledge of the Holocene vegetation history in the coastal plains in Tuscany and Liguria bordering the Mar Ligure (Ligurian sea). In this paper, an assessment of the upper part of the core is presented. During the Holocene, the area was covered by woodlands whose composition changed in time. Numerous oscillations of the tree taxa were recorded which can probably be connected to sea level variations that had a strong influence on the depth of the water table. Human impact is evident particularly at ca. 4200–2500 b.p. when traces of opening of the woodland accompanied by noticeable pollen percentages of Vitis could suggest an early agricultural practice favouring this native plant. The new data have been integrated with previous palaeobotanical and archaeobotanical research. The occurrence of coastal wetlands has been shown. Throughout the Holocene, Mediterranean evergreen vegetation is generally poorly represented; it becomes widespread only in the late Holocene, suggesting that its establishment was probably favoured by the management practices in the coastal plains, particularly land reclamations, starting from the Roman period.     

Vegetation gradients in Atlantic Europe: the use of existing phytosociological data in preliminary investigations on the potential effects of climate change on British vegetation

• 1 This paper aims to demonstrate the use of available vegetation data from the phytosociological literature in preliminary analyses to generate hypotheses regarding vegetation and climate change.
  • 2 Data for over 3000 samples of calcareous grassland, mesotrophic grassland, heath and woodland vegetation were taken from the literature for a region in the west of Atlantic Europe and subjected to ordination by detrended correspondence analysis in order to identify the main gradients present.
    • 3 Climate data were obtained at a resolution of 0.5° from an existing database. The relationship between vegetation composition and climate was investigated by the correlation of the mean scores for the first two ordination axes for each 0.5° cell with the climate and location variables.
      • 4 The ordinations resulted in clear geographical gradients for calcareous grasslands, heaths and woodlands but not for mesotrophic grasslands. Significant correlations were shown between some of the vegetation gradients and the climate variables, with the strongest relationships occurring between the calcareous grassland gradients and July temperature, latitude and oceanicity. Some of the vegetation gradients were also inferred to reflect edaphic factors, management and vegetation history.
        • 5 Those gradients that were related to temperature were hypothesized to reflect the influence of a progressively warmer climate on species composition, providing a baseline for further studies on the influence of climate change on species composition.
        • 6 The validity of the literature data was assessed by the collection of an original set of field data for calcareous grasslands and the subsequent ordination of a dataset containing samples from both the literature and the field. The considerable overlap between the samples from the literature and the field suggest that literature data can be used, despite certain limitations. Such preliminary analyses, using readily available data, can thus achieve useful results, thereby saving lengthy and costly field visits.

     

A synoptic climatology of pollen concentrations during the six warmest months in Sydney,Australia

This paper takes an air mass approach to investigating the influence of weather on pollen concentrations in the atmosphere in Sydney, Australia, by producing a synoptic classification of pollen concentrations measured in the Sydney Basin. This synoptic classification has been produced using multivariate statistical techniques including principal component analysis and cluster analysis, to assign days into meteorologically homogenous categories. Surface and upper air meteorological data for warm months (October-March) over a 10-year period were used as input into the statistical analyses. Eleven synoptic categories were found in Sydney during the warm months. Pollen concentrations for the total pollen load and five individual families measured over a 3.5-year period have been investigated for each of the synoptic categories. High pollen concentrations during the warm months in Sydney are found to be influenced by the presence of a region of low surface pressure located to the south of the continent, bringing fast dry westerly gradient winds to Sydney. It is envisaged that these results will be important from a pollen forecast and associated public health perspective.     

Serpulid-bryozoan-foraminiferal biostromes controlled by temperate climate and reduced salinity: Middle miocene of the Styrian Basin,Austria

Summary Reduced salintiy and a temperate climate prevailed during the Sarmatian in the Styrian Basin, the westernmost embayment of the Central Paratethys. At its northern margin (in the study area) tectonic processes initiated a transgression causing the incision of a cliff into the metamorphic basement and the formation of a carbonate buildup in the latest Sarmatian. The buildup consists of two serpulid-bryozoan-foraminiferal biostromes separated by a microbialite. Serpulids, bryozoa and the encrusting foraminifer Sinzowella caespitosa (Steinmann) as well as microbial mats formed a rigid framework, in which biogenic debris and siliciclastics were baffled. The different biota show complex growth relationships. Stromatolitic crusts and laminated micritic microbial crusts with birdseyes grew in a small sheltered lagoon. The upper biostrome is truncated by a ravinement surface of eustatic orgin and is overlain by ooid grainstone. The buildup drowned during maximum sea-level rise.