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Hydrography and mollusc faunas of the Baltic and the White Sea–North Sea seaway in the Eemian
Palaeogeography, Palaeoclimatology, Palaeoecology 184 (2002) 275^304 www.elsevier.com/locate/palaeo
Hydrography and mollusc faunas of the Baltic and the White Sea^North Sea seaway in the Eemian Svend Funder a; , Igor Demidov b , Yadviga Yelovicheva c a Geological Museum, Ostervoldgade 5^7, DK-1350 Copenhagen K, Denmark Institute of Geology, Karelian Research Centre, Pushkinskaya 11, 185610 Petrozavodsk, Russia Academy of Sciences, Institute of the Geological Sciences of Belarus, Zhodinskaya Street 7, 220141 Minsk, Belarus b
c
Received 28 May 2001; accepted 15 February 2002
Abstract Palynologically dated mollusc and cirriped faunas from a region extending from the North Sea through the Baltic and Karelia to the Arkhangelsk region show that the hydrography of the Baltic was very different from the Holocene. For 2^2.5 ka in the Early Eemian a seaway existed from the Barents to the North Sea through Karelia, until it was severed at the present continental watershed to the north of Lake Onega. After this the Ladoga^Onega trough remained an arm of the Baltic for several millennia. The benthic faunas are comparable to the Holocene, but their boundaries were displaced much further into the Baltic. Notable differences from the Holocene are the absence or rarity of the Macoma balthica biocoenosis, and the presence of cold Portlandia-dominated biocoenoses in Karelia. In the Belt Sea and western Baltic winter sea surface temperatures and salinity were higher than now by ca. 6‡C and 15x, and the distinctly brackish top layer was missing. At the same time cold bottom water (W2.5‡C) with a tendency to anoxia characterised the Karelian arm of the Baltic. Water exchange through this area was inhibited by the constriction to the north of Lake Onega, and the basin and threshold bathymetry. Water transport through the White Sea^Baltic seaway was too sparse to play an active role in the North Atlantic surface circulation or climatic change in the region. The high salinity and temperature in the Belt Sea and western Baltic persisted throughout the Eemian, and are explained by wider and deeper passage from the North Sea to Kattegat, wider straits through Denmark, higher salinity in the North Sea, higher evaporation, as well as more dispersed fresh water supply. The advection of oceanic water into the Baltic culminated in the Early Eemian, before the Carpinus zone, and probably resulted in an oceanic climate in the Baltic region, while at the same time cold winters produced cold bottom water in the Karelian arm. : 2002 Elsevier Science B.V. All rights reserved. Keywords: Baltic; Eemian; molluscs; palynology; interglacial climate; past hydrographical change
1. Introduction During the last interglacial, the Eemian or Mi* Corresponding author. Tel.: +45-35322345; Fax: +45-35322325. E-mail address: [email protected] (S. Funder).
kulinian, the con¢guration and hydrography of the Baltic were very di¡erent from the Holocene, and for some time a seaway connected the White Sea and Barents Sea with the Baltic and North Sea. This paper attempts to assess the geometry, duration, and hydrography of the Baltic in the
0031-0182 / 02 / $ ^ see front matter : 2002 Elsevier Science B.V. All rights reserved. PII: S 0 0 3 1 - 0 1 8 2 ( 0 2 ) 0 0 2 5 6 - 0
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Eemian, and their climatic consequences. This is based on a review of the mollusc faunas and published pollen diagrams from the seaway’s marine sediments (Fig. 1), supplemented with the authors’ own studies on mollusc faunas and a new pollen diagram from a boring in Petrozavodsk. The reconstructed hydrography is based on analogies with present biological and hydrographical conditions in the Baltic and the White Sea summarised in the detailed overviews by Segerstrafile (1957), Voipio (1981), and Rheinheimer (1996) on the Baltic, and similar overviews on the White Sea (Scarlato, 1987; Berger and Naumov, 2000; Galkina et al., 2000), and Zenkevitch (1963) on both areas.
White Sea and the Baltic in the warm Eemian, but the problem remained: while the seaway was warmer than present in both ends, it seemed to be colder than present in the middle (Znamenskaja, 1959; Lavrova, 1962). A large number of Russian contributions have dealt with this problem (e.g. Yakovlev, 1956; Biske, 1959; Lavrova, 1961, 1962; Znamenskaia and Cherminisova, 1962; Biske and Devyatova, 1965; Malakhovskiy et al., 1989; Krasnov et al., 1995), and aspects of this discussion have been reviewed in the west by Raukas (1991) and Donner (1995).
1.1. Previous studies
The Baltic is one of the world’s largest brackish water reservoirs (Fig. 7). This results from high in£ux of fresh water, low evaporation, and restricted in£ow of oceanic water (Kullenberg, 1981). The largest input of fresh water, ca. 40% of the total, comes from rivers around the Gulf of Bothnia and the largest single contributor is the Neva River, which supplies ca. 20% of the total input (Ehlin, 1981). The salt-water input is recruited from surface water in the North Sea and Kattegat (Gustafsson, 1997), but its passage into the Baltic is restricted by the limited capacity of the Danish straits, which transport mainly the out£ow of lighter brackish water. Notable obstacles for the entrance of saline water are the thresholds at Darss and Drogden. The basin and threshold bathymetry of the Baltic further inhibits the progress of the salt water, which forms dense pools with a tendency to anoxia in the basins (Sohlenius et al., 2001). The tidal amplitude is small, and surface circulation and mixing of surface waters are mainly wind-controlled. The large input of fresh water and the restricted in£ow of saline water create a permanent halocline separating an upper layer of brackish (5^10x) water from denser ( 6 20x) bottom waters. At shallow depths the brackish water is characterised by the Macoma balthica biocoenosis (Zenkevitch, 1963), which penetrates north into the northern Gulf of Bothnia and eastwards into the Gulf of Finland until salinities drop below ca. 3.5x (Segerstrafile, 1957). Other frequent species of this biocoenosis
The ‘Cyprina clay’ in Denmark and the sediments of the ‘Boreal Transgression’ of northern Russia have been known to science for more than a century (Forchammer, 1842; Murchison et al., 1845), and by the turn of the century their correlation with the marine Eemian sediments in Holland was recognised in both areas (deGeer, 1896; Madsen et al., 1908). In 1908 a boring in Petrozavodsk showed that marine sediments were also present in the intervening area (Wollosovich, 1908), and later similar sediments were also found in the St. Petersburg area, but the mollusc faunas in these areas were distinctly Arctic and when Zans (1936) and Brander (1937) suggested that the White Sea and the Baltic had indeed been connected by a seaway through Karelia, they correlated it not with the warm Eemian, but with a younger cold period. This correlation was, however, refuted by faunal observations to the north of Lake Onega, which showed that the warm water of the Boreal Transgression had indeed over£own the watershed from the White Sea and invaded northern Karelia (Zemlyakov, 1936; Goretskyi, 1949), and pollen analyses of the marine sediments showed that the ‘Mginskian’, i.e. the marine sediments with cold marine fauna in Karelia and the St. Petersburg area, were indeed contemporaneous with the warm Eemian sea in the western Baltic. Therefore, a seaway did exist between the
1.2. Present hydrography and benthic faunas of the Baltic and the White Sea
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277
Fig. 1. Localities for Eemian marine mollusc assemblages and pollen diagrams used in this paper.
are Mytilus edulis, Cerastoderma glaucum, and Balanus improvisus. In the more salty water of the Belt Sea the M. balthica biocoenosis may on sandy bottoms be replaced by the Abra alba biocoenosis, with Arctica islandica and Corbula gibba as characteristic members (Petersen, 1918; Thorson, 1957). In the deeper water of the southern Baltic where oxygenation is still su⁄cient the benthic fauna is dominated by the Astarte borealis^Macoma calcarea biocoenosis, two species which have their main distribution in the Arctic (Zenkevitch, 1963). Similar to the Baltic, the White Sea is a semienclosed sea with a narrow passage to the ocean. However, water ventilation is more e⁄cient because of the wider entrance and tidal amplitude up to 8 m. The rather warm summers and large input of fresh water creates summer strati¢cation, and surface salinities are generally ca. 25x with temperatures at more than 15‡ (Berger and Nau-
mov, 2000). The cold winters, on the other hand, are responsible for the dense cold water, which forms a stable water mass in all parts of the sea below 25^50 m (Galkina et al., 2000). The large annual temperature variations in the shallow water are restrictive for both warm and cold species, and the fauna is impoverished in comparison with that of the Barents Sea. The shoreface is inhabited by such subarctic species as Mytilus edulis, Macoma balthica, and Semibalanus balanoides with isolated occurrences of the boreal Littorina littorea. The cold bottom water, on the other hand, houses an arctic fauna dominated by Portlandia arctica (Naumov and Fedyakov, 2000). 1.3. Mollusc faunas, methods and terminology The species composition of mollusc and cirriped faunas along the seaway is shown in Table 1, which lists the species in each area together
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POLYPLACOPHORA Lepidochitona cinerea (Linne¤, 1767) Tonicella marmorea (Fabricius, 1780) GASTROPODA Acmaea rubella (Fabricius, 1780) Acmaea virginea (Mu«ller, 1776) Lepeta caeca (Mu«ller, 1776) Puncturella noachina (Linne¤, 1771) Anatoma crispata Fleming, 1828 Gibbula cineraria (Linne¤, 1758) Margarites costalis (Gould, 1841) Margarites groenlandicus (Gmelin, 1794) Margarites helicinus (Phipps, 1774) Margarites vahlii MUller, 1842 Solariella obscura (Couthouy, 1838) Solariella varicosa (Mighels and Adams, 1842) Moelleria costulata MUller, 1842 Bittium reticulatum (da Costa, 1778) Turritella communis (Risso, 1826) Tachyrhynchus erosus (Couthouy, 1838) Littorina littorea (Linne¤, 1758) Littorina compressa Je¡reys, 1865 Littorina obtusata (Linne¤, 1758) Littorina saxatilis (Olivi, 1792) Lacuna parva (da Costa, 1778) Lacuna vincta (Montagu, 1803) Rissoa membranacea (Adams, 1800) Rissoa parva (da Costa, 1779) Rissoa lilacina Re¤cluz, 1843 Alvania je¡reysi (Waller, 1864) Boreocingula castanea Golikov and Kussakin, 1974 Onoba aculeus (Gould, 1841) Onoba semicostata (Montagu, 1803) Pusillina inconspicua (Alder, 1844) Pusillina sarsi (Love¤n, 1846) Caecum glabrum (Montagu, 1803) Hydrobia ulvae (Pennant, 1777) Hydrobia ventrosa (Montagu, 1803) Hyala vitrea (Montagu, 1803) Aporrhais pespelicani (Linne¤, 1758) Velutina velutina (Mu«ller, 1776) Trichotropis borealis (Broderip and Sowerby, 1829) Natica clausa (Broderip and Sowerby, 1829)
s 15 s 15
99.7 72.4 90.7 99.7 100.0 71.3 99.6 80.0 100.0 99.7 99.6 99.7
Arctic Boreal Arctic Arctic Ubiquitous Boreal Arctic Subarctic Ubiquitous Arctic Arctic Arctic
Foreshore/tidal zone Foreshore/tidal zone Shoreface Shoreface Shoreface Shoreface Shoreface Foreshore/tidal zone Shoreface O¡shore Shoreface Shoreface
s 25 s 15 s 15 s 15 s 30 s 15 s 25 s 15 s 25 s 25 s 25 s 25
Arctic Boreal Boreal Arctic Boreal Boreal Subarctic Subarctic Boreal Subarctic Boreal Boreal Boreal Boreal Arctic
Shoreface Foreshore/tidal Shoreface Shoreface Foreshore/tidal Foreshore/tidal Foreshore/tidal Foreshore/tidal Foreshore/tidal Foreshore/tidal Foreshore/tidal Foreshore/tidal Foreshore/tidal O¡shore Foreshore/tidal
s 15 s 15 s 15 s 30 s5 s 30 s5 s5 s 15 s5 s 15 s 25 s 15 s 25 s 30
73.3 73.3 70.5 67.5 60.5 73.3 62.0 60.0 70.0 100 100.0
Boreal Boreal Boreal Boreal Boreal Boreal Boreal Boreal Boreal Ubiquitous Ubiquitous
Foreshore/tidal Shoreface Shoreface Foreshore/tidal O¡shore Foreshore/tidal Foreshore/tidal Shoreface O¡shore Shoreface O¡shore
100.0
Ubiquitous
Shoreface
99.7 65.0 68.0 99.7 73.3 73.3 75.3 75.3 63.0 78.0 62.0 71.3 68.0 73.3 99.7
zone
zone zone zone zone zone zone zone zone zone zone zone
zone zone zone
PALAEO 2866 25-7-02 Cyaan Magenta Geel Zwart
s5 s 25 s5 s 15 s 30 s5 s5 s 30 s 15 s 25 s 30 s 15
Arkhangelsk region
Foreshore/tidal zone Shoreface
Watershed area
Boreal Ubiquitous
Petrozavodsk area
72.4 100.0
St. Petersburg area
Depth
Vistula
Biogeographical characteristics
Belt Sea
Temperature index
Wadden Sea
Species
Salinity tolerance (x)
Table 1 Eemian macrobenthic faunas
F F
F F
F F F F
F
F
F F F F F F F
F F
F
F
F
F
F
F
F
F
F
F
F F F
F
F
F
F F
F
F
F
F
F
F
F
F
F
F
F
F
F F F
F F F
F
F
F F
F
F
F
F
F
F
F
F
F F
F F F
F
F
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279
GASTROPODA Euspira montagui (Forbes, 1838) 71.3 Euspira pallida (Broderip and Sowerby, 100.0 1829) Euspira alderi (Forbes, 1838) 68.0 Amauropsis islandica (Gmelin, 1791) 99.6 Triphora adversa (Montagu, 1803) 62.0 Cerithiopsis barleei (Je¡reys, 1867) 60.0 Cerithiopsis tubercularis (Montagu, 1803) 60.5 Epitonium commune (Lamarck, 1822) 60.5 Epitonium greenlandicum (Perry, 1811) 80.0 Acirsa coarctata (Je¡reys, 1884) 99.7 Trophon clathratus (Linne¤, 1767) 80.0 Trophon truncatus (Stro«m, 1767) 78.0 Nucella lapillus (Linne¤, 1758) 73.7 Buccinum cyaneum Bruguie're, 1792 100.0 Buccinum undatum Linne¤, 1758 80.4 Buccinum undulatum (MUller, 1842) 100.0 Colus holboelli (MUller, 1842) 99.6 Colus islandicus Gmelin, 1791 99.6 Colus sabini (Gray, 1824) 100.0 Neptunea despecta (Linne¤, 1758) 99.6 Turrisipho moebii (Dunker and 71.3 Metzger, 1874) Nassarius reticulatus (Linne¤, 1758) 61.0 Nassarius pygmaeus (Lamarck, 1822) 61.0 Admete viridula (Fabricius, 1780) 100.0 Mangelia brachystoma (Philippi, 1844) 61.0 Oenopota assimilis? (Sars, 1878) 99.7 Oenopota elegans (MUller, 1842) 99.7 Oenopota harpularia (Couthouy, 1839) 99.7 Oenopota nobilis (MUller, 1842) 99.7 Oenopota pyramidalis (Stro«m, 1768) 99.7 Oenopota scalaris abyssicola 99.7 (MUller, 1842) Oenopota trevelliana (Turton, 1834) 82.0 Oenopota turricula (Montagu, 1803) 55.0 Astyris rosacea (Gould, 1840) 99.6 Chrysallida indistincta (Montagu, 1808) 60.5 Chrysallida interstincta (Brown, 1827) 60.5 Chrysallida spiralis (Montagu, 1803) 70.0 Odostomia conoidea (Brocchi, 1814) 63.5 Odostomia eulimoides (Hanley, 1844) 67.5 Odostomia cf. scalaris MacGillivray, 1843 64.0 Odostomia unidentata (Montagu, 1803) 72.4 Turbonilla acuta (Donovan, 1804) 50.0 Turbonilla rufa (Philippi, 1836) 68.0 Turbonilla lactea (Linne¤, 1758) 60.5
Boreal Arctic Boreal Boreal Boreal Boreal Subarctic Arctic Subarctic Subarctic Boreal Ubiquitous Subarctic Ubiquitous Arctic Arctic Ubiquitous Arctic Boreal
Shoreface Shoreface Foreshore/tidal Foreshore/tidal Foreshore/tidal O¡shore O¡shore ? Foreshore/tidal Shoreface Foreshore/tidal Shoreface Shoreface O¡shore Shoreface O¡shore O¡shore Shoreface O¡shore
s 25 s 15 s 15 s 15 s 30 s 15 s 25 ? s 25 s 15 s 25 s 25 s 15 s 30 s 30 s 30 s 30 s 25 s 30
Boreal Boreal Ubiquitous Boreal Arctic Arctic Arctic Arctic Arctic Arctic
Foreshore/tidal Foreshore/tidal Foreshore/tidal Shoreface Shoreface Shoreface Shoreface Foreshore/tidal Shoreface Foreshore/tidal
Subarctic Boreal Arctic Boreal Boreal Boreal Boreal Boreal Boreal Boreal Lusitanic Boreal Boreal
Shoreface Foreshore/tidal Foreshore/tidal Shoreface Foreshore/tidal Foreshore/tidal O¡shore Shoreface Foreshore/tidal Foreshore/tidal Foreshore/tidal O¡shore Foreshore/tidal
zone zone zone
zone zone
zone zone zone
zone zone
zone zone zone zone
zone zone zone zone
PALAEO 2866 25-7-02 Cyaan Magenta Geel Zwart
s 15 s 15 s 25 s 25 s 25 s 25 s 25 s 25 s 25 s 30 s 15 s 15 s 30 s 15 s 15 s 15 s 30 s 30 s 15 s 15 ? s 15 s 30
Arkhangelsk region
s 15 s 15
Watershed area
O¡shore Shoreface
Petrozavodsk area
Boreal Ubiquitous
St. Petersburg area
Depth
Vistula
Biogeographical characteristics
Belt Sea
Temperature index
Wadden Sea
Species
Salinity tolerance (x)
Table 1(Continued).
F F
F F F
F F
F F
F
F F F F F F
F
F
F
F F
F F F F F F
F
F
F
F
F
F
F F
F F F F
F F F F F
F
F
F
F
F F F
F F
?
F F F
F
F
F
280
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Shoreface ? Foreshore/tidal Foreshore/tidal Foreshore/tidal Foreshore/tidal Foreshore/tidal Foreshore/tidal Foreshore/tidal Shoreface Foreshore/tidal Foreshore/tidal
s 15 ? s 15 s 25 s5 s 15 s 30 ? s 15 s 15 s 15 s 15
F
F
F
F
F
F
F
61.0 68.0 61.0 74.0 100.0 99.6
Boreal Boreal Boreal Subarctic Ubiquitous Arctic
Shoreface Shoreface O¡shore O¡shore O¡shore Changing depth requirements Changing depth requirements Shoreface Changing depth requirements O¡shore Changing depth requirements Shoreface Foreshore/tidal zone O¡shore Foreshore/tidal zone Foreshore/tidal zone Foreshore/tidal zone Foreshore/tidal zone Changing depth requirements Foreshore/tidal zone Foreshore/tidal zone Shoreface Foreshore/tidal zone O¡shore Shoreface O¡shore O¡shore Shoreface Shoreface
s 15 s 30 s 15 s 25 s 15 s 15
F
F
Nuculana minuta (Mu«ller, 1776)
80.0
Yoldia hyperborea Torell, 1859 Yoldiella frigida (Torell, 1859)
99.7 100.0
Yoldiella intermedia (Sars, 1865) Yoldiella lenticula (MUller, 1842)
99.7 99.7
Subarctic Arctic Ubiquitous Arctic Arctic
Portlandia arctica (Gray, 1824) 99.7 Portlandia aestuariorum Mossevitch, 1928 99.7 Bathyarca glacialis (Gray, 1824) 99.7 Glycymeris glycymeris (Linne¤, 1758) 62.5 Mytilus edulis Linne, 1758 75.3 Mytilaster lineatus (Gmelin, 1791) 40.0 Mytilaster cf. minimus (Poli, 1795) 48.0 Crenella decussata (Montagu, 1808) 85.1
Arctic Arctic Arctic Boreal Subarctic Lusitanic Lusitanic Subarctic
Musculus discors (Linne¤, 1767) Musculus tumida (Hanley, 1843) Modiolus modiolus (Linne, 1758) Modiolula phaseolina (Philippi, 1844) Pecten maximus (Linne¤, 1758) Aequipecten opercularis (Linne¤, 1758) Arctinula greenlandica (Sowerby, 1842) Chlamys islandica (Mu«ller, 1776) Chlamys varia (Linne¤, 1758) Heteranomia squamula (Linne¤, 1758)
Ubiquitous Boreal Boreal Boreal Boreal Boreal Arctic Subarctic Boreal Subarctic
100.0 70.5 74.9 71.3 67.0 69.0 99.7 85.1 64.0 78.2
zone zone zone zone zone zone zone zone zone
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Arkhangelsk region
Boreal Lusitanic Boreal Boreal Subarctic Boreal Boreal Lusitanic Boreal Ubiquitous Boreal Boreal
Watershed area
Vistula
65.0 40.0 70.0 73.3 80.0 70.0 70.5 53.0 67.0 100.0 68.0 71.3
Petrozavodsk area
Depth
St. Petersburg area
Biogeographical characteristics
Belt Sea
GASTROPODA Ebala nitidissima (Montagu, 1803) Ebala pointeli (de Folin, 1868) Acteon tornatilis (Linne¤, 1758) Diaphana minuta Brown, 1827 Retusa obtusa (Montagu, 1803) Retusa truncatula (Bruguie're, 1792) Retusa umbilicata (Montagu, 1803) Haminoea navicula (da Costa, 1778) Philine aperta (Linne¤, 1767) Cylichna alba (Brown, 1827) Cylichna cylindraceae (Pennant, 1777) Akera bullata (Mu«ller, 1776) BIVALVIA Nucula nitidosa Winckworth, 1930 Nucula nucleus (Linne¤, 1758) Nucula sulcata Bronn, 1831 Nuculoma corticata (MUller, 1842) Nuculoma tenuis (Montagu, 1808) Nuculana pernula Mu«ller, 1779
Temperature index
Wadden Sea
Species
Salinity tolerance (x)
Table 1(Continued).
F F
? F F
F
F
F
F
F
F
F
F
?
F F
F F F F
F
s 15
F
s 15 s 30
F
s 30 s 30
F
s 25 65 s 15 s 30 65 ? ? s 15 s 15 s 15 s 15 s 30 s 15 s 15 s 15 s 25 s 30 s 15
F
F
F
F
F
F
F
F F F
F
F
?
F
F
F
F
F
F F F
F F F
F
F
F
F
F F F F
F
F
F
F F
F
F
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281
Astarte crebricostata Mc Andrews and Forbes, 1847 Astarte crenata (Gray, 1824) Astarte elliptica (Brown, 1827) Astarte montagui montagui Dillwyn, 1817 Astarte montagui striata (Leach, 1819) Astarte sulcata (da Costa, 1778) Acanthocardia echinata (Linne, 1758) Acanthocardia paucicostata (Sowerby, 1841) Parvicardium elegantulum (MUller, 1842) Parvicardium exiguum (Gmelin, 1791) Parvicardium ovale (Sowerby, 1840) Parvicardium scabrum (Philippi, 1844) Plagiocardium papillosum (Poli, 1795) Laevicardium crassum (Gmelin, 1791) Cerastoderma edule (Linne, 1758) Cerastoderma glaucum (Poiret, 1789) Ciliatocardium ciliatum (Fabricius, 1780) Serripes groenlandicus (Bruguie're, 1789) Mactra stultorum (Linne¤, 1758) Spisula elliptica (Brown, 1827) Spisula solida (Linne¤, 1758) Spisula subtruncata (da Costa, 1778) Lutraria lutraria (Linne¤, 1758) Ensis ensis (Linne¤, 1758) Ensis siliqua (Linne¤, 1758) Phaxas pellucidus (Pennant, 1777) Tellina donacina Linne¤, 1758 Tellina fabula Gmelin, 1791 Tellina tenuis da Costa, 1778 Macoma balthica (Linne, 1758) Macoma calcarea (Gmelin, 1791)
Arctic
Shoreface Shoreface Foreshore/tidal zone Shoreface O¡shore Shoreface O¡shore O¡shore O¡shore Foreshore/tidal zone Foreshore/tidal zone Changing depth requirements Shoreface
?
99.6 100.0 85.1 81.0 72.3 72.9 50.0
Arctic Ubiquitous Subarctic Subarctic Boreal Boreal Lusitanic
O¡shore Shoreface Shoreface Shoreface O¡shore Shoreface ?
? s 15 s 15 ? s 30 s 15 ?
73.9 70.0 73.1 70.5 56.0 62.0 71.0 71.0 99.7 99.7 58.0 73.1 57.0 67.5 56.0 62.0 63.5 70.0 55.0 68.5 70.0 85.1 100.0
Boreal Boreal Boreal Boreal Boreal Boreal Boreal Boreal Arctic Arctic Boreal Boreal Boreal Boreal Boreal Boreal Boreal Boreal Boreal Boreal Boreal Subarctic Ubiquitous
O¡shore Shoreface Shoreface Shoreface Shoreface Shoreface Foreshore/tidal zone Foreshore/tidal zone Shoreface Shoreface Foreshore/tidal zone Shoreface Shoreface Foreshore/tidal zone Foreshore/tidal zone Foreshore/tidal zone Foreshore/tidal zone Shoreface O¡shore Foreshore/tidal zone Foreshore/tidal zone Foreshore/tidal zone Changing depth requirements
? s 15 s 15 s 30 ? s 30 s5 65 s 15 s 15 s 30 s 15 s 30 s 15 s 30 s 25 s 25 s 15 s 15 s 30 s 15 65 s5
99.7
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s 15 s 25 s 25 ? s 25 s 25 s 15 s 30 s 25 s 15 s 30 s5
F
Arkhangelsk region
F
Watershed area
Boreal Boreal Lusitanic Lusitanic Boreal Arctic Boreal Boreal Boreal Boreal Boreal Arctic
Petrozavodsk area
70.5 65.5 53.0 51.0 71.3 99.7 62.0 60.5 70.0 70.0 72.0 99.6
Depth
St. Petersburg area
BIVALVIA Pododesmus patelliformis (Linne¤, 1761) Ostrea edulis Linne¤ 1758 Loripes lucinalis (Lamarck, 1818) Lucinella divaricata (Linne¤, 1758) Lucinoma borealis (Linne¤, 1767) Axinopsida orbiculata (Sars, 1878) Thyasira £exuosa (Montagu, 1803) Hemilepton nitidum (Turton, 1822) Montacuta ferruginosa (Montagu, 1808) Mysella bidentata (Montagu, 1803) Turtonia minuta (Fabricius, 1780) Astarte borealis (Schumacher, 1817)
Vistula
Biogeographical characteristics
Belt Sea
Temperature index
Wadden Sea
Species
Salinity tolerance (x)
Table 1(Continued).
F
F
F
F
F
F
F F F
F
F
F
F
F
F
F
F
F
F F F
F
F
F
F F
F
F
F
F
F F F
F
F
?
F
F
F
F
F
F F
F
F
F
F F F F
F
F F
F
F
F
F
F
F
F F
F F F
F
? F
F
F
F
F F
F
F
F F F F F
F
F
F
F
F F
F
F
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Balanus improvisus Darwin, 1854 Semibalanus balanoides (Linne¤, 1758)
69.0 80.4
Foreshore/tidal zone Shoreface Shoreface Changing depth requirements Foreshore/tidal zone Foreshore/tidal zone
s 30 s5 s5 s 25
F
65 s 15
F
Boreal Subarctic
with information on their requirements in relation to temperature, salinity and depth, as described below. With few exceptions the faunas come from shoreface habitats and re£ect surface temperature and salinity. The taxonomical nomenclature generally follows Le Renard (1997) and
F
F
F
F
F
F
F
Arkhangelsk region
Boreal Ubiquitous Subarctic Subarctic
F
Watershed area
s 15 s 30 s5 s5 s 30 s 25 65 s5 s 30 s 25 s 25 s 30 s 25 s 30 s 30 ? s 15 s 15 s5 s 15 s5 s 25 s 15 s 15 s5 s 25 s 15 s 25 s 25 s 15
Petrozavodsk area
Shoreface Shoreface Foreshore/tidal zone Shoreface O¡shore O¡shore Shoreface Foreshore/tidal zone O¡shore Shoreface O¡shore Shoreface O¡shore Shoreface Shoreface ? Shoreface O¡shore Shoreface O¡shore Shoreface Shoreface O¡shore Shoreface Shoreface O¡shore Shoreface O¡shore Foreshore/tidal zone Shoreface
St. Petersburg area
Lusitanic Boreal Boreal Boreal Boreal Boreal Lusitanic Boreal Boreal Boreal Boreal Boreal Boreal Boreal Boreal ? Boreal Boreal Ubiquitous Boreal Ubiquitous Boreal Boreal Boreal Boreal Boreal Arctic Boreal Boreal Boreal
Vistula
Depth
Belt Sea
BIVALVIA Gastrana fragilis (Linne¤, 1758) 48.0 Donax vittatus (da Costa, 1778) 58.5 Scrobicularia plana (da Costa, 1778) 62.0 Abra alba (Wood, 1802) 67.5 Abra nitida (Mu«ller, 1776) 70.5 Abra prismatica (Montagu, 1803) 70.5 Abra segmentum (Re¤cluz, 1843) 47.0 Arctica islandica (Linne, 1767) 72.4 Kelliella abyssicola (Forbes, 1844) 69.0 Chamelea gallina (Linne¤, 1757) 71.3 Timoclea ovata (Pennant, 1777) 71.3 Gouldia minima (Montagu, 1803) 60.0 Dosinia lupinus (Linne¤, 1758) 70.0 Tapes decussata (Linne¤, 1758) 62.0 Paphia aurea (Gmelin, 1791) 63.5 Paphia senescens (Cocconi, 1873) 0.0 Venerupis senegalensis (Montagu, 1803) 70.0 Mysia undata (Pennant, 1777) 67.5 Mya truncata Linne¤, 1758 100.0 Corbula gibba (Olivi, 1792) 70.5 Hiatella arctica (Linne¤, 1767) 100.0 Panomya arctica (Spengler, 1793) 73.3 Saxicavella je¡reysi Winckworth, 1930 68.0 Barnea candida (Linne¤, 1758) 64.0 Zirfaea crispata (Linne, 1767) 71.3 Thracia convexa (Wood, 1815) 60.5 Thracia myopsis (MUller, 1842) 99.7 Thracia papyracea (Poli, 1791) 68.0 Thracia villosiuscula (Macgillivray, 1827) 60.0 Verruca stroemia (Mu«ller, 1776) 73.3 CIRRIPEDIA Chtamalus stellatus (Poli, 1791) 55.0 Balanus balanus (Linne¤, 1758) 100.0 Balanus crenatus (Bruguie're, 1789) 75.3 Balanus hameri (Ascanius, 1767) 85.1
Biogeographical characteristics
Wadden Sea
Temperature index
Species
Salinity tolerance (x)
Table 1 (Continued).
F
F
F
F F
F
F
F
F
F
F
F
F
F F
F
F
F
F
F
F
F
F
F F F
F
F
F F
F F
F
F
F
F
F
F F
F F
F
F
F
F
F
F
F F
F F F F
F
F
F F
F F
F
F
F
F F
Poppe and Goto (1991, 1993), who give comprehensive synonyms. 1.4. Biogeographical zones The species are classi¢ed according to their
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283
Fig. 2. Distribution of marine biogeographical zones in the North Atlantic. (A) Present, (B) Early Eemian.
northern penetration along the European coasts (Fig. 2). The biogeographical terms are here used in an oceanographic sense, based on water mass characteristics, and deviate slightly from Feyling-Hanssen’s (1955) often-used scheme, which was based on the occurrence of indicator species along the Norwegian coast. The Lusitanian zone is characterised by dominance of warm Atlantic water as seen especially in the high winter sea surface temperatures, s 9‡C, and summer temperatures higher than 16‡. The boundary for this zone, which marks the northernmost occurrence of a number of Mediterranean species, is at the southwestern entrance to the English Channel (Ekman, 1953). A notable representative of this fauna is Lucinella divaricata, which penetrated far into the Baltic during the Eemian. Also the Boreal zone is strongly in£uenced by warm Atlantic surface water. Summer sea surface temperatures are higher than 8‡C and the area is year-round ice-free. This zone is bounded to the north and east by the limit for regular occurrence of winter sea ice at the Murman coast. The zone extends for almost 2000 km from S to N along the North European coast, and is characterised throughout by its uniform shallow water faunas with Arctica islandica, Spisula elliptica, Cerastoderma edule, and Zirfaea crispata.
During the Eemian this zone extended through the Barents and Kara Seas to Taymyr (Fig. 2). To the north and east of this, the subarctic zone is the zone of mixing of Polar and Atlantic water. Summer sea surface temperatures are higher than 3‡C and although winter sea ice is present, there is at least a 2-month ice-free period. The foreshore and intertidal fauna (e.g. Mytilus edulis, Semibalanus balanoides, Macoma balthica) has its northern limit in this area. Further to the north and east cold polar water and long lasting ice cover characterise the arctic zone. The long lasting ice cover makes stationary life on the foreshore di⁄cult and the foreshore fauna is missing. The upper shoreface is inhabited by such species as Portlandia arctica, Arctinula greenlandica, and Bathyarca glacialis. The cold polar water from the Arctic basin sinks down and occurs as deep water far to the south. The arctic species may therefore occur at greater depths even in the boreal zone. Finally, some species have a wide tolerance towards temperature and are ubiquitous in the entire region. 1.5. Temperature As an expression of temperature demands the lusitanian, boreal, and subarctic species are in-
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dexed after their northern boundaries (Table 1). Along the south^north trending European coastline this is expressed as latitude (72.6 = 72‡60PN). Along the west^east trending Russian arctic coastline it has been adjusted by calibration with species with a northern limit on Svalbard. The arctic species are indexed according to their southern boundaries (90.72Wsouthern limit at 72‡N). The distribution and temperature demands are based on information from Jensen and Spa«rck (1934), Stephensen (1933, 1938), Gayevskaja (1948), Filatova (1957), Tarasov and Zevina (1957), Zenkevitch (1963), Tebble (1966), Scarlato (1981, 1987), HUisZter (1986), Graham (1988), Seaward (1990, 1993), Golikov (1995), and Bratlegaard and Holthe (1997). 1.6. Salinity Life in water with reduced salinity is di⁄cult for most marine organisms, and the lower the salinity the lower the number of species that can endure it. Also, reduced in£ow of oceanic water means stronger in£uence of atmospheric temperatures and larger annual temperature variations, making life di⁄cult for both warmth and cold demanding species. In Table 1 the species are grouped after their tolerance to reduced salinity into euhaline ( s 30x), and polyhaline ( s 25x), b-mesohaline ( s 15x), a-mesohaline ( s 5x), and oligohaline ( 6 5x). The species’ salinity demands are based on information from Scarlato (1981, 1987) and Golikov (1995), and their occurrence on the salinity gradient from the open ocean through the North Sea and into the Baltic (Seaward, 1990, 1993; Jensen and Knudsen, 1995; KUie et al., 2000). Common brackish-tolerant species are Mytilus edulis, Macoma balthica, and Balanus improvisus. In the Arctic Portlandia aestuariorum inhabits, as the only mollusc, estuaries with salinities down to less than 1x (Scarlato, 1981; Petryashov et al., 1999). Most low salinity-tolerant species can thrive in oceanic water and estimating salinity depends on the species with the least tolerance to brackish conditions. Among the most demanding oceanic representatives can be mentioned Mactra stultorum and Tapes decussatus.
1.7. Depth The benthic species are not dependent on water depth. However, since such environmental parameters as wave action, temperature variation, and photosynthesis are related to depth this is a major determining factor for their distribution. In the shallow water wave action and tide are probably the most important restrictive factors, and the species are here categorised according to their shallowest depth on the shore plane from foreshore/tidal to shoreface and o¡shore. Many arctic molluscs prefer to live high on the shoreface in the Arctic, but extend their distribution to o¡shore environments in the boreal zone, and therefore have changing depth requirements. The depth parameter is used here to indicate habitat diversity in each area. 1.8. Age We date and correlate the Eemian marine sediments by using published pollen diagrams from marine sediments, supplemented with a new diagram from Petrozavodsk. The Eemian in northern Europe is characterised by a uniform development of vegetation, and similar pollen zones can be identi¢ed in the entire area of deciduous forest (Zagwijn, 1996), which extended from the Atlantic coasts eastwards to Moscow and to Lake Onega in the north (Grichuk, 1984). In the Early Eemian the forest trees immigrated very rapidly and seem to have arrived everywhere in the same order of succession. Since the whole immigration period was short ( 6 3 ka), the di¡erence in arrival time of a species to di¡erent parts of the region may not have amounted to more than a few centuries (Zagwijn, 1996; Cheddadi et al., 1998). After the immigration rush came the period of stability, beginning with the Carpinus zone. In this period the di¡erences in timing of pollen zones are probably much greater, and it would be unlikely if the period of deciduous forest in the north of Russia was not considerably shorter than in the south, whereas the subsequent taiga stages, the Picea and Pinus zones, were longer. Since we do not know the migration routes for the trees, this error cannot be estimated, but it
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may not a¡ect our results seriously, because the events discussed here mainly took place in the Early Eemian. The durations of the pollen zones in Table 2 are Mu«ller’s (1974) estimates, derived from the counting of annual layers in lake sediments from North Germany. The duration of the early pollen zones, the ¢rst 2.1 ka, was done with high precision, but the duration of the remaining part of the Eemian had to be made by extrapolation and is much more uncertain (Mu«ller, 1974). Tentatively we suggest absolute ages for the pollen zones (Table 2). This is based on a correlation of Zagwijn’s (1983) pollen-dated sea level curve from the southern North Sea with curves for eustatic sea level change based on U^Th dating of corals (Stirling et al., 1998; Esat et al., 1999; McCulloch et al., 1999; Henderson and Slowey, 2000). The correlation ¢x-point in the two records is the attainment of the Eemian sea level highstand, here estimated as ca. 500 yr after the beginning of the Carpinus zone and dated on coral reefs to 128 [ 1 ka (McCulloch and Esat, 2000; Shackleton, 2000). From this we arrive at tentative ages for the Eemian pollen zones in northern Europe (Table 2)1 . This estimate is tentative and ignores isostatic e¡ects, which could have terminated the transgression in the area before the attainment of the global highstand, and shifted the beginning of the ‘pollen-Eemian’ further back in time if isostatic uplift was faster than sea level rise; or the transgression could have lasted longer if the area was subjected to forebulge collapse. Disregarding these factors may to some extent be justi¢ed by sea level curves (Esat et al., 1999; McCulloch et al., 1999; Israelson and Wohlfahrt, 1999), which indicate that sea level rise was so rapid that it is likely to have overridden the isostatic e¡ects, and the change from very high to zero rate of sea level rise was
1
This estimate has the consequence that the ‘pollen-Eemian’ here ends at 120.5 ka, and not at the well established age of ca. 115 ka. It is beyond the scope of this paper to discuss the cause for this discrepancy. However, it should be noted that the duration of the Late Eemian pollen zones is based on extrapolation, which could have minimised their durations (Mu«ller, 1974).
285
abrupt, suggesting that maximum transgression was coeval with the attainment of the global highstand. This is supported by the fact that regression in many areas within the region began in the Carpinus zone.
2. Description of areas Table 1 gives lists of molluscs and cirripeds from each area along the seaway shown on Fig. 1. Notes are given below of their type of occurrence, pollen analytical results, and outline of marine inundation. The duration of marine inundation at each site is shown in Table 2. In some areas, notably the Gulf of Bothnia, there are no mollusc records, and the marine environment is deduced from diatoms. 2.1. The Wadden Sea This area comprises the southern North Sea, southwestern Jutland and northwestern Germany. Eemian marine sediments are known from numerous borings along the 130 km coast, which lies outside the margin of Weichselian glaciation. The sedimentary successions comprise basal sandy shoreface facies, ‘senescens sand’, named after Paphia senescens, overlying limnic sediments, and overlaid by thick ‘Turritella clay’, named after Turritella communis, or ‘Olander beds’, which are strati¢ed ¢ne sand and clay, probably tidal (Dittmer, 1951; Temmler, 1995). Further inland the sediments show great variability and re£ect a wide variety in hydrographic conditions in this area of narrow straits and broads (Gripp, 1964; Kosack and Lange, 1985; Temmler, 1995). In general the successions show a transition from limnic to brackish to marine to brackish to limnic, and re£ect a complete transgressive^regressive cycle. The mollusc fauna in Table 1 represents all these sedimentary facies and is based on the studies of Heck and Brockmann (1950), Dittmer (1951, 1954), Jaeckel (1963), Hinsch (1985), and especially Nordmann (1928). The rich fauna owes partly to the diversity in habitats and hydrography, but also to the high salinity.
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Table 2 Correlation and duration of pollen zones along the White Sea^Baltic seaway
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PALAEO 2866 25-7-02 Cyaan Magenta Geel Zwart Dark shading marks maximum duration of White Sea^Baltic seaway. (1) Based on the listed durations and correlation between pollen- and U^Th-dated sea level curves (Zagwijn, 1983; Esat et al., 1999; Henderson and Slowey, 2000).
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In Schleswig-Holstein the transgression came in the Betula^Pinus zone (Menke, 1985), as evidenced by drowning surfaces and transgression conglomerates with Ostrea edulis showing that both sea surface temperatures and salinity were high at the very beginning of the Eemian (Hinsch, 1985; Knudsen, 1985; Meijer and Preece, 1995). This implies the English Channel was open at the beginning of the Eemian and transported more Atlantic water into the North Sea than during the Holocene (Meijer and Preece, 1995). An even warmer and more saline fauna is known from Holland, but not pollen analytically dated within the Eemian (Spaink, 1958; Meijer and Preece, 1995). The large proportion of oceanic species, which do not appear in the North Sea at present (e.g. Cerithiopsis tubercularis, Odostomia conoidea, Tapes decussatus), show that maximum salinity, s 30x, was reached in the Corylus, Taxus/Tilia, and early Carpinus zone. After this, regression set in and during the Picea zone the sea retreated from this area. The Eemian sea inundated valleys and depressions and gave the coastline a complex outline with small irregular inlets penetrating deep into the country and land areas extending out into what is now the southern North Sea, maybe as far as Helgoland (Dittmer, 1951, 1954; Gripp, 1964; Behre, 1970; Konradi, 1976; Kubisch and Scho«nfeld, 1985; Knudsen, 1994; Friborg, 1996). In the area of the Kiel Canal in situ marine sediments outline a winding sound with broads and narrows extending across Jutland from the Friesian islands eastwards to areas behind the Weichselian ice margin, less than 20 km from the present Baltic coast. This made Kosack and Lange (1985) suggest the existence of a seaway from the North Sea to the Baltic. 2.2. Kattegat and the Belt Sea The inner Danish coasts and those of northern Germany as far as Mecklenburg^Vorpommern were overridden by ice during the Weichselian, and almost all occurrences of marine Eemian sediments are dislocated clasts. The fauna list in Table 1 is compiled from Nordmann (1928),
287
Heck and Brockmann (1950), Gehl (1961), Madsen (1965), and Houmark-Nielsen (1994). The Cyprina clay, named after Cyprina (now Arctica) islandica, is exposed in coastal cli¡s around the western Baltic in Denmark and northern Germany, possibly as far east as Ru«gen and Usedom (Konradi, 1976; Kubisch and Scho«nfeld, 1985; Ho«£e et al., 1985; Mu«ller et al., 1995; Ru«hberg et al., 1995). The lower contact is usually a drowning surface overlying limnic sand or clay. At its top the Cyprina clay is usually truncated by glacial or glaci£uvial sediments, only in the cli¡s at Gammelmarke does the clay grade into shoreface sand rich in shells, the ‘Tapes sand’, named after Tapes (now Paphia) senescens with a fauna similar to that in the senescens sand in Schleswig-Holstein. Dislocated clasts containing a deep-water fauna similar to the Turritella clay of Schleswig-Holstein are also known from a few sites (\dum, 1933; Houmark-Nielsen, 1994). The faunas contain oceanic species, which nowadays are absent from the North Sea and Kattegat (Turbonilla lactea, Gouldia minima, Hemilepton nitidum) or they live in the North Sea and northern Kattegat, but not in southern Kattegat and the Belt Sea (Caecum glabrum, Mactra stultorum). All these oceanic species are, however, known only from the Gammelmarke cli¡s in the westernmost Baltic (Madsen et al., 1908), but in the eastern Belt Sea polyhaline species such as Ostrea edulis, Turritella communis, and Aporrhais pespelicani show that salinities here also were higher than those attained during the Holocene and the common occurrence of Lucinella divaricata shows that temperatures were higher. This shows that the Belt Sea, after a brief brackish phase, had salinities of 25^30x. At the western edge of the basin the salinity was at least periodically even higher. At Ristinge the marine inundation has recently been dated to the beginning of the Quercus zone, and the marine inundation lasted ca. 3.4 ka, until the early Carpinus zone (Kristensen et al., 2000). Ostracodes, foraminifera and dino£agellates indicate an initial brackish phase followed shortly afterwards by more saline conditions, which culminated at the start of the Carpinus zone
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(Table 2), although fully marine conditions were never attained (Head and Gibbard, 2000). Also in eastern Schleswig-Holstein and Mecklenburg the marine inundation took place before the Carpinus zone and the retreat came in the Picea zone (Heck and Brockmann, 1950; Gehl, 1961).
higher than 25x (Fig. 3). By the end of the Carpinus zone salinity dropped, as evidenced by the low diverse Mytilus edulis^Cerastoderma glaucum assemblages.
2.3. Western Baltic
In the Neva lowland and areas around Lake Ladoga Eemian marine sediments consist of dark organic and gas rich laminated clay and silt with thin layers of ¢ne sand, known both from numerous borings and from temporary exposures in clay pits (MalakhovskiY et al., 1989; Krasnov et al., 1995). The sediments occur both in situ and as glacially dislocated clasts. They represent o¡shore or lagoonal mud deposited as basin ¢ll in drowned valleys and depressions (Znamenskaia and Cherminisova, 1962; Malakhovskiy et al., 1989). The best-known occurrence is at Mga. Pollen diagrams from this and nearby Rybatskoe show that the marine drowning began early in the Eemian, in the Pinus^Ulmus zone, and the marine phase lasted until the Pinus zone (Table 2 and Znamenskaja, 1959; Lavrova, 1962). The mollusc faunas from Mga and other sites have been described by Skorokhod (1932) and Lavrova (1961, 1962), and are preserved in the Geological Museum in St. Petersburg where we have had the opportunity to study them. With a total of only eight species the faunas are notable for their low diversity and lack of warm species. They are characterised especially by Portlandia arctica and Portlandia aestuariorum (Skorokhod, 1932). The fauna shows increasing and then decreasing salinity upwards in the sediments with maximum salinity being achieved prior to the Corylus zone (Fig. 3). This is shown by the transition from predominance of P. arctica in the low-
During the Eemian the Vistula valley was inundated by the sea up to 70 km from its present mouth in the Danzig Bay. The sediments, considered to be in situ, are known from rare exposures and many borings and comprise shoreface sand and o¡shore clay (Makowska, 1986, 1991). The mollusc faunas in Table 1 have been compiled from a number of sites (Brodniewics, 1960, 1969; Makowska, 1986), especially Elblag and Nowiny, which display normal transgressive^regressive sequences. The domination of Corbula gibba throughout indicates a protected fjord environment. In the Nowiny boring the marine episode began at the onset of the Quercus zone and lasted until some time into the Picea zone (Makowska, 1986). Paphia senescens was among the very ¢rst immigrants, and Lucinella divaricata followed immediately after, showing that the connection to the North Sea was established and sea surface temperatures already were higher than Holocene. Retusa umbilicata and Abra nitida are sparsely present throughout, and indicate that salinity culminated in the Quercus zone, periodically rising to over 30x. The poor development of Ostrea and Venerupis senegalensis (earlier pullastra, Brodniewics, 1960, 1969), on the other hand, shows that conditions were not optimal for them, suggesting that salinity probably generally was not
2.4. Eastern Baltic
Fig. 3. The Baltic^White Sea region at the peak of marine inundation in the Early Eemian, tentatively dated to the period 132^ 130 ka. Red arrows mark entranceways discussed in the text. The development of salinity at each locality is shown in columns at the base of the ¢gure. The columns represent the duration of the Eemian divided into pollen zones. The outline of the seaway is based on an assessment of maps in: Apukhtin and Krasnov, 1966; Atlas Karelskoi ASSR, 1989; Biske, 1959; Biske and Devyatova, 1965; Devyatova, 1961, 1982; Devyatova and Loseva, 1964; Forsstro«m et al., 1988; Friborg, 1996; Gehl, 1961; Goretskyi, 1949; Gripp, 1964; Ho«£e et al., 1985; International Quaternary map of Europe, 1970^1971; Knudsen, 1994; Konradi, 1976; Kosack and Lange, 1985; Lavrova, 1960, 1961; Legkova and Schukin, 1967; Lundqvist and Robertsson, 1994; Makowska, 1986; Malakhovskiy et al., 1989; Ru«hberg et al., 1995; Seidenkrantz et al., 2000; Temmler, 1995; Znamenskaia and Cherminisova, 1962. Bathymetry is modelled from present depths and Seidenkrantz and Knudsen (1997).
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Fig. 4. Smear of Portlandia aestuariorum in laminated clayey silt from Mga (from collections in the Geological Museum, St. Petersburg).
er part to predominance of the more brackish P. aestuariorum from the beginning of the Carpinus zone (Skorokhod, 1932). Portlandia is present from the earliest phase (Lavrova, 1962) and is concentrated in smears and thin lenses, probably indicating mass death and gentle in-habitat reworking (Fig. 4). The other components of the fauna occur as single individuals scattered in the sediment. They are present only at the salinity maximum in the middle part of the sequence and comprise Macoma calcarea, thin-shelled small Mytilus edulis and a few Cerastoderma, originally identi¢ed as C. edule, but their triangular appearance makes it more likely that they belonged to the more brackish Cerastoderma glaucum. These shells, together with leaf fragments of Zostera marina, were probably redeposited from the nearby foreshore (Lavrova, 1962). M. calcarea, C. glaucum and a single worn Littorina littorea, as well as the small and thin M. edulis suggest surface salinity at ca. 15x when it peaked in the middle sequence. Portlandia arctica, on the other hand, generally requires salinity of more than 25x (e.g. Mossevitch, 1928; Scarlato, 1987), suggesting strati¢cation between low saline surface water and denser bottom water. We suggest that the high-saline and warm planktonic diatoms (Brander, 1937; Znamenskaia and Cherminisova, 1962) could represent brief injections of warm saline water surface water from the Baltic, such as those that control the salinity of the Baltic at present (Bo«rngen et al., 1990). The scarcity of molluscs, their concentration in
smears or in sand laminae, and the high organic content of the sediments indicate that anoxic bottom conditions were frequent, but interrupted by short intervals of oxygenation. Anoxic conditions would also explain the rarity of benthic and attached diatoms, which induced Znamenskaia and Cherminisova (1962) to assume a water depth of 80^100 m over the sites. At Prangli, at the mouth of the Gulf of Finland, an Eemian pollen diagram from marine sediments from a boring shows a gradual change from Late Saalian limnic and brackish phase and into ‘normal marine’ conditions in the Early Eemian (Liivrand, 1991). A sudden change from brackish to marine took place by the end of the Pinus^Ulmus zone. At this place in the core mollusc shells also appear to have their ¢rst appearance, which we consider to mark the marine inundation. The marine conditions continue into the early Picea zone, and are ended by a hiatus (Fig. 3). These results have recently been con¢rmed and expanded by pollen^foraminifer- and diatom analyses of Eemian marine sediments in three borings in coastal Latvia and the Riga Bay (Kalnina, 2001). In the boring at Plasumi the marine sediments cover the entire Eemian, beginning and ending with cold brackish water facies in the Late Saalian and Early Weichselian. Diatoms and foraminifera indicate that water depth and salinity, s 18x based on diatoms, culminated before the Carpinus zone, when regression set in (Kalnina, 2001). On the eastern shore of Lake Ladoga marine
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reworked megaclasts of Eemian marine sediments have been encountered in borings and pollen diagrams give minimum ages for the duration of the marine inundation of this basin, lasting from the Quercus and into the early Picea zone (Devyatova, 1982). 2.5. The Gulf of Bothnia In situ marine sediments in borings at three localities in Ostrobothnia were deposited immediately after deglaciation or overlie lacustrine sediments during fall in relative sea level (Eriksson et al., 1980, 1999). Pollen diagrams and correlation with sea level history indicate that the marine inundation began after the immigration of Quercus and lasted at least until within the Picea zone. Initially, high percentages of polyhalobous diatoms indicate salinities higher than 17x. Upwards in the sediments a decline in salinity began before the onset of the Carpinus zone (Fig. 3). In a boring at Norra Sanna«s in central Sweden marine Eemian clay is topped by clay with fresh water diatoms, and the marine inundation lasted from the Quercus zone until some time before the beginning of the Carpinus zone (Robertsson et al., 1997). Reworked brackish-marine sediments occur also at other sites and indicate that salinity in the Early Eemian was higher than Holocene and sea level fall began before the immigration of Picea (Robertsson and Garc|¤a Ambrosiana, 1992; Robertsson, 2000). 2.6. Karelia, Petrozavodsk Marine Eemian sediments are known from borings in isolated sediment pockets preserved in depressions of the Precambrian basement. In the city of Petrozavodsk borings have shown marine Eemian sediments considered to be in situ at 40 m above sea level (asl). They show continuous development from glaciolacustrine clay to marine sediments and lacustrine sand and till (Wollosovich, 1908; Lukashov, 1982). The lower clay contains plant remains and is dominated by Macoma calcarea, while the upper sandier facies is bioturbated and dominated by Portlandia aestuariorum and thin-shelled Mytilus edulis. The mollusc fauna
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suggests salinities at 10^15x, based on M. calcarea, at the base to somewhat lower at the top. The bioturbation shows that anoxia was not prevalent as in the St. Petersburg area (Wollosovich, 1908; Skorokhod, 1932). Pollen diagrams from two borings in the city of Petrozavodsk contain su⁄cient pollen of deciduous trees for the record to be correlated with the West European pollen stratigraphy (Fig. 5 and Lukashov, 1982). These results show that the marine inundation took place in the Early Eemian, in the Pinus^Ulmus zone. In the diagram presented here the marine episode lasted until the end of the Corylus zone (Fig. 5), while the earlier diagram by Devyatova (in Lukashov, 1982) sets the boundary somewhat later in the upper part of the following Taxus^Tilia zone. By palynological correlation this corresponds to a duration of ca. 2.5 ka, tentatively from ca. 132.4 to ca. 130 ka. Petrozavodsk is close to the continental watershed towards the White Sea, and the sedimentary sequence can be considered as a monitor for marine over£ow from the White Sea, implying that the over£ow and the seaway existed only in the Early Eemian. After this the connection was severed to the north of Lake Onega, but the Ladoga^ Onega Lake basin probably continued as a brackish arm of the Baltic for some time. The outline of the Eemian basin in Fig. 3 is reconstructed using evidence from Apukhtin and Krasnov (1966), Atlas Karelskoi ASSR (1989), and Devyatova (1961, 1982). The two lake basins were connected by a wide shallow sound, 50 km across at the narrowest, and with scattered islands. In its northern end the Onega basin opened up towards the White Sea. 2.7. The continental watershed area Between Lake Onega and the White Sea shore 150 km to the north. The marine Eemian sediments occur as isolated patches ¢lling up depressions in the bedrock surface (Zemlyakov, 1936; Goretskyi, 1949; Biske, 1959; Biske and Devyatova, 1965; Heikki et al., 1995). The mollusc fauna listed in Table 1 is derived from borings on the south side of the watershed (Povenets) and from a number of isolated occur-
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PALAEO 2866 25-7-02 Cyaan Magenta Geel Zwart Fig. 5. Pollen diagram from boring in the city of Petrozavodsk, period of maximum marine inundation and inferred existence of the White Sea^Baltic seaway shaded. Pollen frequencies calculated according to Grichuk (1961).
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rences on the watershed up to altitudes of 96^101 m (Goretskyi, 1949; Biske, 1959; Devyatova and Loseva, 1964). With 24 species the faunas are signi¢cantly richer than those from nearby Petrozavodsk and St. Petersburg. Astarte sulcata, Trichotropis borealis, and Nucella lapillus show that warm saline Atlantic water from the White Sea £owed over the threshold and into the Onega basin. This is con¢rmed by the presence of Panomya arctica and N. lapillus at Povenets and saline diatoms at nearby Medvezhegorsk on the Onega side of the watershed. Also here the marine interval seems to be restricted to within the period from the Quercus^Pinus to the Taxus/Tilia zones (Znamenskaia and Cherminisova, 1962; Biske and Devyatova, 1965). During the maximum transgression the watershed located immediately to the north of Lake Onega at 105 m asl was penetrated by two 30^ 40-km-long and ca. 10-km-wide sounds (Fig. 1), which opened northwards into a wide shallow archipelago extending to the present White Sea shore (Atlas Karelskoi ASSR, 1989; Goretskyi, 1949; Apukhtin and Krasnov, 1966). This may be somewhat conjectural because the present watershed area is composed of Weichselian sediments. However, the faunal contrast between areas on and to the north of the watershed and those to its south show that during the Eemian a major hydrographical constriction of the seaway existed at this place. 2.8. Arkhangelsk region The Severnaya Dvina basin is the classical area for sediments from the Boreal Transgression. The basin contained a shallow marine inlet up to 500 km from the present river mouth at the White Sea. In the coastal area this basin was, via its tributary valleys, fused with the basins of the neighbouring Onega and Mezen rivers to form a large shallow archipelago extending from the Onega River basin to the Timan Ridge (Devyatova, 1982). Most of the area was ice-covered during the Weichselian (Larsen et al., 1999), but the Eemian sediments are mostly undisturbed and in situ. They are exposed in riverbanks and comprise
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every conceivable coastal and o¡shore habitat. Reviews of the very comprehensive literature dealing with the Eemian in this area were given by Lavrova (1937), Legkova (1967), and Devyatova (1982). In the coastal area to the north of Arkhangelsk pollen analysis from a boring at Izhma shows that the marine transgression reached the present coastline of the White Sea in the Late Saalian and marine conditions persisted into the Weichselian (Pleshivtseva, 1972). On the very southern margin of the basin, at Pasva, the marine inundation was restricted to the Quercus zone (Devyatova, 1982). Midway between these two localities, at Krasnaya Gorka, the transgression began in the Quercus zone and lasted into the Picea zone (Fig. 3 and Devyatova, 1982). The Eemian marine sediments begin with a lower shoreface facies. This is bounded below by a £ooding or ravinement surface, and followed by up to 30-m-thick o¡shore mud; followed again by shoreface and £uvial sediments. Further upriver shoreface sand dominates the sequences (Lavrova, 1937). The fauna listed in Table 1 is based on the lists by Zharkhidze (1963), Legkova and Schukin (1967), and our own observations, and comprises a wide range of habitats. This is to some extent the explanation for the unusual richness with 112 species, the richest among the regions presented here. The earliest transgression sediments contain a high number of oceanic species, such as Boreocingula castanea and Astarte sulcata as well as several Yoldiella species, showing that during this early phase even the most remote areas were oceanic or polyhaline. A transition to Macoma balthica-dominated faunas in the upper shoreface sediments at many localities may indicate that a change to estuarine conditions with reduced salinity took place at the start of the regression after the Quercus and Quercus^Corylus zones, as suggested by Devyatova (1982).
3. Geometry and bathymetry of the seaway The seaway was, according to our reconstruction, a ca. 150-km-wide sound extending from the
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North Sea to the south and west coast of the present White Sea, and via a broad connection further north to the Barents Sea, a distance of ca. 2500 km (Fig. 3). During the Holocene the three Danish straits, LillebZlt, StorebZlt and \resund, are the restrictive factor for water exchange between the Baltic and the ocean. In their present state these straits were shaped as £uvial valleys, eroded by rivers that drained the Baltic during its fresh water stages in the Late Weichselian and Early Holocene (Bjo«rck, 1995; Andre¤n et al., 2000). StorebZlt today carries 75% of the brackish outlet water from the Baltic and almost all in£ux of oceanic water from Kattegat (e.g. Segerstrafile, 1957), but there is no evidence to indicate that it existed in the Eemian. The same can be said for the narrow winding LillebZlt. Along \resund and in the interior of SjZlland, on the other hand, typical Eemian molluscs (Lucinella divaricata, Paphia senescens) occur scattered in Weichselian meltwater sediments and beach sediments (\dum, 1933; Madsen, 1965). Although sparse and circumstantial, this could indicate that a major entranceway to the Baltic traversed SjZlland. A candidate for such a waterway could be the buried Alnarp^Esrum Valley. This is a more than 100-km-long, more than 10-km-wide and more than 50-mdeep rami¢ed valley system crossing southern Sweden and northern SjZlland on its way from Kattegat to the Baltic (Miller, 1977; Konradi, 1992; Houmark-Nielsen, 1999). The valley system was shaped by faulting in the Palaeogene and Mesozoic substrate, and last functioned as a meltwater channel during the Weichselian when it was ¢lled with sand and gravel (Houmark-Nielsen, 1999). Although no Eemian sediments occur in the valley, the scattered ¢nds of reworked Eemian shells in its vicinity could indicate that it served as a major corridor for water exchange between the Baltic and Kattegat during the Eemian, thereby circumventing the thresholds at Darss and Drogden, which presently form a barrier for the entrance of dense oceanic water into the Baltic. As noted above a connection from the ocean to the Baltic may also have existed through Schleswig-Holstein in the area of the Kiel Canal, but the
outline of this waterway, a winding pattern of broads and narrow passages (Kosack and Lange, 1985), indicates that it was similar to the present Limfjord traversing northern Jutland without playing a signi¢cant role in the water exchange of the Baltic. In the north, from a much-enlarged White Sea the entrance gradually narrowed southwards over northern Karelia towards the major constriction at the watershed north of Lake Onega (Goretskyi, 1949; Atlas Karelskoi ASSR, 1989). The dimension of this entrance was in its dimensions similar to the present step-by-step entrance to the Baltic from the North Sea through Kattegat and the Danish straits, and both its outline and the faunal contrast between the areas to the north and south suggest that this was a major obstacle limiting the capacity for water transport through the seaway. A shallow side entrance from the Severnaya Dvina basin into Karelia may have existed along the river Voda (Devyatova, 1982). A peculiar feature of the Eemian coastline is its lobate appearance with the sea everywhere forming saline estuaries in present day valleys and depressions, contrasting to the present mature coastlines. This suggests rapid sea level rise and/or high tidal amplitude. Since this feature is found in all parts of the region rapid sea level rise may be the most likely explanation. The bathymetry of the seaway on Fig. 3 is based mainly on present depth contours to which are added the maximum altitude of the Eemian marine inundation, but no correction has been made for glacial erosion during the Weichselian when both the Baltic and the White Sea served major outlets for the Scandinavian ice sheet. As at present the Baltic basin was composed of a series of deep troughs with depths exceeding 200 m, separated by shallower thresholds. The same pattern was probably found in the Karelian passage where deep troughs occurred in the Ladoga and Onega depressions. The most signi¢cant deviation from the present bathymetry is the trough that may have extended across northern Jutland and into Kattegat, as indicated by borings showing great thickness of marine sediments of probable Eemian age (Lykke-Andersen et al., 1993; Seidenkrantz and Knudsen, 1997). The trough, which
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Table 3 Biogeographical and salinity grouping in Eemian faunas
apparently was ¢lled up during the Eemian, marks a signi¢cant deviation from the Holocene Kattegat, and may have played an important role in the hydrography. In summary, the White Sea^Baltic seaway was a ca. 150-km-wide sound extending for ca. 2500 km from the North Sea to the southern Barents Sea composed of more than 200-m-deep basins, separated by shallow thresholds. A major constriction existed in the north, to the north of Lake Onega, while a deep trench in Kattegat and a deep valley system extending through SjZlland and Skafine may have a¡orded a more e⁄cient water exchange than the Holocene. A channel through Schleswig-Holstein probably supplied restricted in£ow from the North Sea.
4. Duration The history of the seaway is a product of isostasy and eustasy following removal of the Saalian ice sheet, and at each site the duration of sea cover is determined by the interaction between these two forces. Since the palynological results come from the coastal fringes of the seaway, they can give only a minimum estimate of the duration of sea cover in each area.
The premises for pollen analytical dating of the marine sediments in each area have been discussed above, and the duration of marine inundation in each area is shown in Table 2 and Fig. 3. Outside the seaway, at the entranceways on the shore of the White Sea and in Kattegat borings have shown continuous sea cover from the Late Saalian through the Eemian and into the Weichselian (Pleshivtseva, 1972; Knudsen, 1994; Seidenkrantz and Knudsen, 1997; Kristensen et al., 1998). In the Wadden Sea marine invasion with warm fauna took place already in the Betula^Pinus zone, at the beginning of the Eemian. Recent results from Latvia indicate that the Baltic was also inundated with saline water at this time (Kalnina, 2001). Shortly afterwards, in the Pinus^Ulmus zone ca. 100 yr into the Eemian, Petrozavodsk was inundated, showing that the seaway into the Baltic was now opened to the Barents Sea. In£ux from the North Sea is documented ¢rst in the faunas of the Vistula valley from the beginning of the Quercus zone, 200 yr later. At this time also the northern parts of the Gulf of Bothnia were probably inundated. These results could indicate that the seaway opened ¢rst to the north and shortly afterwards to the west. However, as noted above, the pollen zones are not precisely
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Fig. 6. Inferred development of salinity in the southern Baltic during the Holocene and the Eemian. Salinity colours as in Fig. 3. The Holocene development is based on results from the Bornholm basin (Sohlenius et al., 2001).
synchronous in this large region, and are not suf¢ciently well de¢ned to distinguish events which are only a few hundred years apart, but it can be concluded that the opening of the Baltic to the Barents Sea and to the North Sea came in close succession within the ¢rst centuries of the Eemian. The regression began in the Corylus or Taxus/ Tilia zone (Petrozavodsk, Gulf of Bothnia). At many sites there is evidence for shallowing of the water depth in the Carpinus zone, and in the Picea zone the water retreated from many areas (Schleswig-Holstein, Vistula, Prangli). The records from Mga and the Ladoga shore show that that the Baltic was still in connection with the ocean at the end of the Eemian. From this, the Baltic was in continuous connection with the ocean at least from the beginning to the end of the Eemian, and there were, at least in the southern Baltic, no fresh water stages like the
Early Holocene Ancylus stage. The seaway from the North Sea to the White Sea functioned only in the Early Eemian from the Pinus^Ulmus or the Quercus zone to within the Taxus/Tilia zone, a period of 2^2.5 ka.
5. Hydrography and climate Table 3 summarises information on distribution, salinity and number of species recorded in each area. 5.1. Salinity and temperature In the Belt Sea the Cyprina clay contains a sparse fauna dominated by such small and short-lived species as Corbula gibba. In its structure the fauna is similar to the Abra alba bio-
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coenosis, which also today inhabits these areas (Thorson, 1957). This indicates that also in the Eemian this was an area of protected fjords with shallow wave base, large in£ux of terrestrial nutrients, and reduced salinity, as shown also by the frequent occurrence of Balanus improvisus. However, at the few localities where it has been preserved the shoreface Tapes or senescens sand may with its frequent Chamaelea gallina and Spisula elliptica compare to the C. gallina biocoenosis, which today inhabits sandy bottoms in the more saline Kattegat, but is absent from the Belt Sea and the Baltic (Thorson, 1957). The fauna in the scattered occurrences of Turritella clay is very similar to today’s Amphiura biocoenosis on deeper water in Kattegat (Thorson, 1957). This indicates that during the Eemian the salinity both in the upper and lower part of the water column in the Belt Sea and the western Baltic was as high as those of the present Kattegat, ca. 25x. This is supported by the absence or rarity of the Macoma balthica biocoenosis, which has inhabited the Baltic shallow water during the Holocene (Segerstrafile, 1957), and the presence of Ostrea and Venerupis senegalensis as well as other salinity demanding species in the Vistula valley indicating a surface salinity of ca. 25x, which is 15x higher than now and ca. 10x higher than the Holocene maximum. The evidence from Poland indicates that this salinity lasted from the Quercus zone and until regression set in at the end of the Carpinus zone (Makowska, 1986). In addition, the occurrence of Lusitanian species, notably Lucinella divaricata, in this area indicates winter sea surface temperatures at ca. 9‡C, 6‡C warmer than now. Fig. 6 compares the inferred salinity development during the Holocene and the Eemian in the southern Baltic. The Holocene is deduced from a record from the Bornholm basin, characterised by its short Ancylus stage and long and brackish initial Littorina stage (Sohlenius et al., 2001). The Early Eemian history was apparently simpler than that of the Holocene and lacks the repeated changes between fresh and brackish. We ascribe this to the dominance of eustatic sea level rise over isostatic rebound in the Eemian, as discussed above. Apart from this di¡erence the de-
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velopment seems to be in phase, with the salinity maximum being achieved in the Early Holocene and Eemian ^ only the Eemian Baltic was considerably more saline throughout. We explain this by a combination of factors as discussed below. In the White Sea area and Arkhangelsk region the situation was similar to the southern Baltic. The shoreface faunas, which now are subarctic, were dominated by such boreal species as Arctica islandica, Spisula elliptica and Cerastoderma edule, showing that salinities and temperatures were considerably higher than now, and the area was probably year-round ice-free. This lasted at least until the Carpinus zone, when regression began. In contrast, the low diverse faunas of Karelia and the St. Petersburg area, seven to eight species, re£ect as noted above low salinity and temperature and, in the St. Petersburg area, probably widespread anoxia. The dominating species, Portlandia arctica, is stenothermal and Jensen (1942) found that it requires bottom water temperatures below 4‡C, and preferably below 2.5‡C. Somewhat higher surface temperatures are indicated by the occurrence of Cerastoderma glaucum and Mytilus edulis. The Portlandia fauna has no analogue in the present Baltic, but similar biocoenoses exist in isolated bays with restricted water exchange along the coasts of the White Sea (Zenkevitch, 1963). In these bays the shallow waters are heated to high summer temperatures, as much as 20‡, and the shoreface fauna is composed of M. edulis, Macoma balthica and such boreal outliers as Littorina littorea and the plant Zostera marina, while the poorly oxygenated denser bottom water at 15 m depth with 6 0‡ temperatures houses an arctic P. arctica biocoenosis. During the Eemian the poor benthic faunas in the Karelian passage therefore point to long lasting water mass strati¢cation with low saline (ca. 15x) surface water and denser ( s 25x) cold (W2.5‡) bottom water with frequent anoxia. It should be noted that although P. arctica nowhere occurs as far south as St. Petersburg, the present faunas in the Baltic do have an arctic aspect, especially among crustaceans, as discussed intensively for more than a century (e.g. Segerstrafile, 1957), and Astarte borealis and Macoma calcarea, two species with their main distribution
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Fig. 7. Inferred surface hydrography of the Baltic at present (A) and Eemian (B). Colours for surface salinity as in Fig. 3. Arrows: red: warm surface current; blue: cold bottom current; green: brackish surface current (W10x); striped red/yellow: coastal water surface current ( s 15x); yellow: major source of river runo¡. Thickness of arrow signi¢es quantity. Present hydrography based on Segerstrafile, 1957; Zenkevitch, 1963; Kullenberg, 1981; Ehlin, 1981.
in the Arctic, thrive in the cold bottom water of the southern Baltic (Zenkevitch, 1963). The lack of other arctic species could possibly be a consequence of the Ancylus Lake stage, which in the Early Holocene interrupted the marine history of the Baltic (Fig. 6 and Bjo«rck, 1995; Andre¤n et al., 2000). Uninterrupted marine conditions going back to the Late Saalian may be a part of the explanation for the persistence of the Portlandia fauna at these southerly latitudes. 5.2. Hydrography and climate Fig. 7 compares the Early Eemian hydrography with that of the present. Conspicuous di¡erences are the larger size of the eastern Baltic, and the higher surface salinity of the southern Baltic in the Eemian. During the Holocene the restricted capacity of the Danish straits has been the main restriction for in£ow of saline water into the Baltic, and we suggest that the much higher salinity during the Eemian was partly a result of wider and deeper entranceways through Denmark. As
noted above the entrance between Norway and Denmark was wider, and a deep trench may have existed in Kattegat. Further, the Alnarp^Esrom valley could possibly have provided a short cut between Kattegat and the Baltic, and a more limited in£ow may have occurred through Schleswig-Holstein (Fig. 3). Other factors also played an important role. Thus the salinity in the recruiting area, the North Sea, was higher, as shown by the faunas in the Wadden Sea; and higher temperatures caused higher evaporation from the sea surface. Better mixing of the surface water, as indicated by the apparent absence of the Macoma balthica biocoenosis, may have been a¡orded by wind as at present and/or more vigorous tidal currents. Finally, the fresh water input was more dispersed. This applies especially to the Neva drainage, which today delivers ca. 20% of the Baltic fresh water input concentrated into the Gulf of Finland (Ehlin, 1981). In the Eemian this supply was distributed over and possibly caught in the Karelian basins. As noted above the faunas in Karelia and the
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St. Petersburg area suggest strati¢cation between low saline surface water and denser and colder bottom water with frequent anoxia. This implies very restricted advection of saline water into the Karelian passage and cold winters in this area, but during the period of the White Sea^Baltic seaway there was at least occasionally over£ow of saline surface water from the White Sea into the Lake Onega basin. This is shown by the saline fauna at Povenets. However, the contrast between this and the fauna from Petrozavodsk and the St. Petersburg area indicates that the saline in£ow both from the north and the south was insu⁄cient for ventilating the Karelian basin. In the St. Petersburg area the warm and saline £ora of planktonic diatoms may also imply £uxes of advective surface water from the southern Baltic, but it seems likely that a major restriction to the £ow must have existed also in the area of the Gulf of Finland. These results show that the White Sea^Baltic seaway during its 2^2.5-ka existence was not a major corridor for north^south transport of cold or warm water, and it is unlikely that its opening and closure had any signi¢cant in£uence on the north European climate. This is in contradiction to theoretical hypotheses expressed recently in a number of papers to explain changing oceanic and continental climates in northern Europe during the Eemian (Zagwijn, 1996; Kristensen et al., 1998; Bjo«rck et al., 2000; Rioual et al., 2001). Instead we suggest that these di¡erences may ¢nd their cause in the larger extent of the Eemian Baltic, and its more vigorous ventilation, both of which culminated in the Early Eemian, and gradually changed to regression and lower salinity.
6. Conclusions Our studies suggest that ^ the Baltic was connected with the ocean throughout the Eemian. In the Belt Sea and southern Baltic temperature and salinity were generally ca. 6‡C warmer and 15x higher than at present, culminating in the Early Eemian before the beginning of the Carpinus zone. After this the sea regressed and salinity declined.
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^ a seaway through Karelia connected the North Sea with the Barents Sea for 2^2.5 ka until isostatic uplift severed it at the watershed to the north of Lake Onega during the Taxus/Tilia zone or earlier. The Onega^Ladoga region continued to be an arm of the Baltic for several millennia, the Ladoga arm probably until the end of the Picea zone. ^ the seaway never functioned as a corridor for large-scale transport of water masses, and its opening and closure probably did not e¡ect the general ocean circulation or north European climate. ^ the higher salinity and more vigorous circulation in the Baltic and Belt Sea were caused by larger capacity of the Danish entranceways, higher salinity in the recruiting area of the North Sea, higher evaporation, and a more dispersed supply of river runo¡. ^ the passage between the present Baltic and White Sea, Karelia and the St. Petersburg region, had long lasting strati¢cation with brackish surface water and dense cold bottom water and frequent anoxia, similar to present conditions in the Gulf of Bothnia. ^ the contrast between the warm and well ventilated western and southern Baltic and the cold and stagnant water of Karelia and St. Petersburg implies that the Eemian oceanic to continental climate gradient from west to east over northern Europe was steeper than the Holocene.
Acknowledgements This study was carried out under the auspices of the ESF (European Science Foundation) administered QUEEN project (Quaternary environments, Eurasian Arctic), and the EU (European Union) Balteem project (Palaeoenvironmental and palaeoclimatic evolution of the Baltic Sea basin during the last (Eemian, Mikulino) interglacial). Fieldwork in Russia and Denmark was funded by the Danish Research Council’s grant to the CATLINA and CLIENT projects. All these projects provided fora for numerous and valuable discussions. We are especially indebted to Valery Astakhov, Dmitri Malakovskiy, and
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Alexei Matiouchkov for help and guidance. The artwork was performed by Lisa Bellhage, Louise Hansen, and Bent Knudsen. Very valuable comments and stimulating discussion were provided by Tom Meijer, and the two reviewers Matti Eronen and Richard Preece.
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Wollosovich, K.A., 1908. Petrozavodskiy morskoy postpliotsen (in Russian). Mater. Geol. Russl. 23, 298^318. Yakovlev, S.A., 1956. Osnovy geologii chetvertichnykh otlozheniy Russkoi ravniny. Stratigraphia (The fundamentals of the Quaternary geology of the Russian Plain. Stratigraphy). Moscow, 314 pp. (in Russian). Zagwijn, W.H., 1983. Sea-level changes in the Netherlands during the Eemian. Geol. Mijnb. 62, 437^450. Zagwijn, W.H., 1996. An analysis of Eemian climate in western and central Europe. Quat. Sci. Rev. 15, 451^469. Zans, V., 1936. Das Letztinterglaziale Portlandia-Meer des Baltikums (in German). Bull. Comm. Ge¤ol. Finl. 115, 231^ 250. Zemlyakov, B.F., 1936. Chetvertichnaia geologia Karelii. Karelskiy Nauchno-Issledovatelskiy Institut, Trudy Sektsii Estestvennykh Proizvoditelnykh sil 1, 102 pp. (in Russian). Zenkevitch, L., 1963. Biology of the Seas of the USSR. Allen and Unwin, London, 955 pp.
Zharkhidze, V.S., 1963. K istorii razvitiya iogo-vostochnoy chasti Barentseva morya i ego fauny s verkhnechetvertichnogo vremeni. Ukhtinskoye territorialnoe geologichyeskoye upravdyenie, Moskovskiy gosudarstvyennyy universitet M.V. Lomonosova, izdatelstvo Moskovskogo universiteta, pp. 91^98 (in Russian). Znamenskaja, O.M., 1959. Stratigra¢cheskoe polozhenie Mginskikh morskikh otlozheniy (in Russian). Dokl. Akad. Nauk SSSR 129, 401^404. Znamenskaia, O.M., Cherminisova, Ye.A., 1962. Rasprostranyeniye Mginskogo mezhlednikovykh morya i osnovnyie cherty ego paleogeogra¢i. In: Lavrova, M.A., Fadeyeva, A.P., Zhingarev-Dobroselsky, A.T., (Eds.), Problems of Stratigraphy of the Quaternary in the North-West European USSR, pp. 125^139 (in Russian).
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Hydrography and mollusc faunas of the Baltic and the White Sea^North Sea seaway in the Eemian Svend Funder a; , Igor Demidov b , Yadviga Yelovicheva c a Geological Museum, Ostervoldgade 5^7, DK-1350 Copenhagen K, Denmark Institute of Geology, Karelian Research Centre, Pushkinskaya 11, 185610 Petrozavodsk, Russia Academy of Sciences, Institute of the Geological Sciences of Belarus, Zhodinskaya Street 7, 220141 Minsk, Belarus b
c
Received 28 May 2001; accepted 15 February 2002
Abstract Palynologically dated mollusc and cirriped faunas from a region extending from the North Sea through the Baltic and Karelia to the Arkhangelsk region show that the hydrography of the Baltic was very different from the Holocene. For 2^2.5 ka in the Early Eemian a seaway existed from the Barents to the North Sea through Karelia, until it was severed at the present continental watershed to the north of Lake Onega. After this the Ladoga^Onega trough remained an arm of the Baltic for several millennia. The benthic faunas are comparable to the Holocene, but their boundaries were displaced much further into the Baltic. Notable differences from the Holocene are the absence or rarity of the Macoma balthica biocoenosis, and the presence of cold Portlandia-dominated biocoenoses in Karelia. In the Belt Sea and western Baltic winter sea surface temperatures and salinity were higher than now by ca. 6‡C and 15x, and the distinctly brackish top layer was missing. At the same time cold bottom water (W2.5‡C) with a tendency to anoxia characterised the Karelian arm of the Baltic. Water exchange through this area was inhibited by the constriction to the north of Lake Onega, and the basin and threshold bathymetry. Water transport through the White Sea^Baltic seaway was too sparse to play an active role in the North Atlantic surface circulation or climatic change in the region. The high salinity and temperature in the Belt Sea and western Baltic persisted throughout the Eemian, and are explained by wider and deeper passage from the North Sea to Kattegat, wider straits through Denmark, higher salinity in the North Sea, higher evaporation, as well as more dispersed fresh water supply. The advection of oceanic water into the Baltic culminated in the Early Eemian, before the Carpinus zone, and probably resulted in an oceanic climate in the Baltic region, while at the same time cold winters produced cold bottom water in the Karelian arm. : 2002 Elsevier Science B.V. All rights reserved. Keywords: Baltic; Eemian; molluscs; palynology; interglacial climate; past hydrographical change
1. Introduction During the last interglacial, the Eemian or Mi* Corresponding author. Tel.: +45-35322345; Fax: +45-35322325. E-mail address: [email protected] (S. Funder).
kulinian, the con¢guration and hydrography of the Baltic were very di¡erent from the Holocene, and for some time a seaway connected the White Sea and Barents Sea with the Baltic and North Sea. This paper attempts to assess the geometry, duration, and hydrography of the Baltic in the
0031-0182 / 02 / $ ^ see front matter : 2002 Elsevier Science B.V. All rights reserved. PII: S 0 0 3 1 - 0 1 8 2 ( 0 2 ) 0 0 2 5 6 - 0
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Eemian, and their climatic consequences. This is based on a review of the mollusc faunas and published pollen diagrams from the seaway’s marine sediments (Fig. 1), supplemented with the authors’ own studies on mollusc faunas and a new pollen diagram from a boring in Petrozavodsk. The reconstructed hydrography is based on analogies with present biological and hydrographical conditions in the Baltic and the White Sea summarised in the detailed overviews by Segerstrafile (1957), Voipio (1981), and Rheinheimer (1996) on the Baltic, and similar overviews on the White Sea (Scarlato, 1987; Berger and Naumov, 2000; Galkina et al., 2000), and Zenkevitch (1963) on both areas.
White Sea and the Baltic in the warm Eemian, but the problem remained: while the seaway was warmer than present in both ends, it seemed to be colder than present in the middle (Znamenskaja, 1959; Lavrova, 1962). A large number of Russian contributions have dealt with this problem (e.g. Yakovlev, 1956; Biske, 1959; Lavrova, 1961, 1962; Znamenskaia and Cherminisova, 1962; Biske and Devyatova, 1965; Malakhovskiy et al., 1989; Krasnov et al., 1995), and aspects of this discussion have been reviewed in the west by Raukas (1991) and Donner (1995).
1.1. Previous studies
The Baltic is one of the world’s largest brackish water reservoirs (Fig. 7). This results from high in£ux of fresh water, low evaporation, and restricted in£ow of oceanic water (Kullenberg, 1981). The largest input of fresh water, ca. 40% of the total, comes from rivers around the Gulf of Bothnia and the largest single contributor is the Neva River, which supplies ca. 20% of the total input (Ehlin, 1981). The salt-water input is recruited from surface water in the North Sea and Kattegat (Gustafsson, 1997), but its passage into the Baltic is restricted by the limited capacity of the Danish straits, which transport mainly the out£ow of lighter brackish water. Notable obstacles for the entrance of saline water are the thresholds at Darss and Drogden. The basin and threshold bathymetry of the Baltic further inhibits the progress of the salt water, which forms dense pools with a tendency to anoxia in the basins (Sohlenius et al., 2001). The tidal amplitude is small, and surface circulation and mixing of surface waters are mainly wind-controlled. The large input of fresh water and the restricted in£ow of saline water create a permanent halocline separating an upper layer of brackish (5^10x) water from denser ( 6 20x) bottom waters. At shallow depths the brackish water is characterised by the Macoma balthica biocoenosis (Zenkevitch, 1963), which penetrates north into the northern Gulf of Bothnia and eastwards into the Gulf of Finland until salinities drop below ca. 3.5x (Segerstrafile, 1957). Other frequent species of this biocoenosis
The ‘Cyprina clay’ in Denmark and the sediments of the ‘Boreal Transgression’ of northern Russia have been known to science for more than a century (Forchammer, 1842; Murchison et al., 1845), and by the turn of the century their correlation with the marine Eemian sediments in Holland was recognised in both areas (deGeer, 1896; Madsen et al., 1908). In 1908 a boring in Petrozavodsk showed that marine sediments were also present in the intervening area (Wollosovich, 1908), and later similar sediments were also found in the St. Petersburg area, but the mollusc faunas in these areas were distinctly Arctic and when Zans (1936) and Brander (1937) suggested that the White Sea and the Baltic had indeed been connected by a seaway through Karelia, they correlated it not with the warm Eemian, but with a younger cold period. This correlation was, however, refuted by faunal observations to the north of Lake Onega, which showed that the warm water of the Boreal Transgression had indeed over£own the watershed from the White Sea and invaded northern Karelia (Zemlyakov, 1936; Goretskyi, 1949), and pollen analyses of the marine sediments showed that the ‘Mginskian’, i.e. the marine sediments with cold marine fauna in Karelia and the St. Petersburg area, were indeed contemporaneous with the warm Eemian sea in the western Baltic. Therefore, a seaway did exist between the
1.2. Present hydrography and benthic faunas of the Baltic and the White Sea
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277
Fig. 1. Localities for Eemian marine mollusc assemblages and pollen diagrams used in this paper.
are Mytilus edulis, Cerastoderma glaucum, and Balanus improvisus. In the more salty water of the Belt Sea the M. balthica biocoenosis may on sandy bottoms be replaced by the Abra alba biocoenosis, with Arctica islandica and Corbula gibba as characteristic members (Petersen, 1918; Thorson, 1957). In the deeper water of the southern Baltic where oxygenation is still su⁄cient the benthic fauna is dominated by the Astarte borealis^Macoma calcarea biocoenosis, two species which have their main distribution in the Arctic (Zenkevitch, 1963). Similar to the Baltic, the White Sea is a semienclosed sea with a narrow passage to the ocean. However, water ventilation is more e⁄cient because of the wider entrance and tidal amplitude up to 8 m. The rather warm summers and large input of fresh water creates summer strati¢cation, and surface salinities are generally ca. 25x with temperatures at more than 15‡ (Berger and Nau-
mov, 2000). The cold winters, on the other hand, are responsible for the dense cold water, which forms a stable water mass in all parts of the sea below 25^50 m (Galkina et al., 2000). The large annual temperature variations in the shallow water are restrictive for both warm and cold species, and the fauna is impoverished in comparison with that of the Barents Sea. The shoreface is inhabited by such subarctic species as Mytilus edulis, Macoma balthica, and Semibalanus balanoides with isolated occurrences of the boreal Littorina littorea. The cold bottom water, on the other hand, houses an arctic fauna dominated by Portlandia arctica (Naumov and Fedyakov, 2000). 1.3. Mollusc faunas, methods and terminology The species composition of mollusc and cirriped faunas along the seaway is shown in Table 1, which lists the species in each area together
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POLYPLACOPHORA Lepidochitona cinerea (Linne¤, 1767) Tonicella marmorea (Fabricius, 1780) GASTROPODA Acmaea rubella (Fabricius, 1780) Acmaea virginea (Mu«ller, 1776) Lepeta caeca (Mu«ller, 1776) Puncturella noachina (Linne¤, 1771) Anatoma crispata Fleming, 1828 Gibbula cineraria (Linne¤, 1758) Margarites costalis (Gould, 1841) Margarites groenlandicus (Gmelin, 1794) Margarites helicinus (Phipps, 1774) Margarites vahlii MUller, 1842 Solariella obscura (Couthouy, 1838) Solariella varicosa (Mighels and Adams, 1842) Moelleria costulata MUller, 1842 Bittium reticulatum (da Costa, 1778) Turritella communis (Risso, 1826) Tachyrhynchus erosus (Couthouy, 1838) Littorina littorea (Linne¤, 1758) Littorina compressa Je¡reys, 1865 Littorina obtusata (Linne¤, 1758) Littorina saxatilis (Olivi, 1792) Lacuna parva (da Costa, 1778) Lacuna vincta (Montagu, 1803) Rissoa membranacea (Adams, 1800) Rissoa parva (da Costa, 1779) Rissoa lilacina Re¤cluz, 1843 Alvania je¡reysi (Waller, 1864) Boreocingula castanea Golikov and Kussakin, 1974 Onoba aculeus (Gould, 1841) Onoba semicostata (Montagu, 1803) Pusillina inconspicua (Alder, 1844) Pusillina sarsi (Love¤n, 1846) Caecum glabrum (Montagu, 1803) Hydrobia ulvae (Pennant, 1777) Hydrobia ventrosa (Montagu, 1803) Hyala vitrea (Montagu, 1803) Aporrhais pespelicani (Linne¤, 1758) Velutina velutina (Mu«ller, 1776) Trichotropis borealis (Broderip and Sowerby, 1829) Natica clausa (Broderip and Sowerby, 1829)
s 15 s 15
99.7 72.4 90.7 99.7 100.0 71.3 99.6 80.0 100.0 99.7 99.6 99.7
Arctic Boreal Arctic Arctic Ubiquitous Boreal Arctic Subarctic Ubiquitous Arctic Arctic Arctic
Foreshore/tidal zone Foreshore/tidal zone Shoreface Shoreface Shoreface Shoreface Shoreface Foreshore/tidal zone Shoreface O¡shore Shoreface Shoreface
s 25 s 15 s 15 s 15 s 30 s 15 s 25 s 15 s 25 s 25 s 25 s 25
Arctic Boreal Boreal Arctic Boreal Boreal Subarctic Subarctic Boreal Subarctic Boreal Boreal Boreal Boreal Arctic
Shoreface Foreshore/tidal Shoreface Shoreface Foreshore/tidal Foreshore/tidal Foreshore/tidal Foreshore/tidal Foreshore/tidal Foreshore/tidal Foreshore/tidal Foreshore/tidal Foreshore/tidal O¡shore Foreshore/tidal
s 15 s 15 s 15 s 30 s5 s 30 s5 s5 s 15 s5 s 15 s 25 s 15 s 25 s 30
73.3 73.3 70.5 67.5 60.5 73.3 62.0 60.0 70.0 100 100.0
Boreal Boreal Boreal Boreal Boreal Boreal Boreal Boreal Boreal Ubiquitous Ubiquitous
Foreshore/tidal Shoreface Shoreface Foreshore/tidal O¡shore Foreshore/tidal Foreshore/tidal Shoreface O¡shore Shoreface O¡shore
100.0
Ubiquitous
Shoreface
99.7 65.0 68.0 99.7 73.3 73.3 75.3 75.3 63.0 78.0 62.0 71.3 68.0 73.3 99.7
zone
zone zone zone zone zone zone zone zone zone zone zone
zone zone zone
PALAEO 2866 25-7-02 Cyaan Magenta Geel Zwart
s5 s 25 s5 s 15 s 30 s5 s5 s 30 s 15 s 25 s 30 s 15
Arkhangelsk region
Foreshore/tidal zone Shoreface
Watershed area
Boreal Ubiquitous
Petrozavodsk area
72.4 100.0
St. Petersburg area
Depth
Vistula
Biogeographical characteristics
Belt Sea
Temperature index
Wadden Sea
Species
Salinity tolerance (x)
Table 1 Eemian macrobenthic faunas
F F
F F
F F F F
F
F
F F F F F F F
F F
F
F
F
F
F
F
F
F
F
F
F F F
F
F
F
F F
F
F
F
F
F
F
F
F
F
F
F
F
F F F
F F F
F
F
F F
F
F
F
F
F
F
F
F
F F
F F F
F
F
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279
GASTROPODA Euspira montagui (Forbes, 1838) 71.3 Euspira pallida (Broderip and Sowerby, 100.0 1829) Euspira alderi (Forbes, 1838) 68.0 Amauropsis islandica (Gmelin, 1791) 99.6 Triphora adversa (Montagu, 1803) 62.0 Cerithiopsis barleei (Je¡reys, 1867) 60.0 Cerithiopsis tubercularis (Montagu, 1803) 60.5 Epitonium commune (Lamarck, 1822) 60.5 Epitonium greenlandicum (Perry, 1811) 80.0 Acirsa coarctata (Je¡reys, 1884) 99.7 Trophon clathratus (Linne¤, 1767) 80.0 Trophon truncatus (Stro«m, 1767) 78.0 Nucella lapillus (Linne¤, 1758) 73.7 Buccinum cyaneum Bruguie're, 1792 100.0 Buccinum undatum Linne¤, 1758 80.4 Buccinum undulatum (MUller, 1842) 100.0 Colus holboelli (MUller, 1842) 99.6 Colus islandicus Gmelin, 1791 99.6 Colus sabini (Gray, 1824) 100.0 Neptunea despecta (Linne¤, 1758) 99.6 Turrisipho moebii (Dunker and 71.3 Metzger, 1874) Nassarius reticulatus (Linne¤, 1758) 61.0 Nassarius pygmaeus (Lamarck, 1822) 61.0 Admete viridula (Fabricius, 1780) 100.0 Mangelia brachystoma (Philippi, 1844) 61.0 Oenopota assimilis? (Sars, 1878) 99.7 Oenopota elegans (MUller, 1842) 99.7 Oenopota harpularia (Couthouy, 1839) 99.7 Oenopota nobilis (MUller, 1842) 99.7 Oenopota pyramidalis (Stro«m, 1768) 99.7 Oenopota scalaris abyssicola 99.7 (MUller, 1842) Oenopota trevelliana (Turton, 1834) 82.0 Oenopota turricula (Montagu, 1803) 55.0 Astyris rosacea (Gould, 1840) 99.6 Chrysallida indistincta (Montagu, 1808) 60.5 Chrysallida interstincta (Brown, 1827) 60.5 Chrysallida spiralis (Montagu, 1803) 70.0 Odostomia conoidea (Brocchi, 1814) 63.5 Odostomia eulimoides (Hanley, 1844) 67.5 Odostomia cf. scalaris MacGillivray, 1843 64.0 Odostomia unidentata (Montagu, 1803) 72.4 Turbonilla acuta (Donovan, 1804) 50.0 Turbonilla rufa (Philippi, 1836) 68.0 Turbonilla lactea (Linne¤, 1758) 60.5
Boreal Arctic Boreal Boreal Boreal Boreal Subarctic Arctic Subarctic Subarctic Boreal Ubiquitous Subarctic Ubiquitous Arctic Arctic Ubiquitous Arctic Boreal
Shoreface Shoreface Foreshore/tidal Foreshore/tidal Foreshore/tidal O¡shore O¡shore ? Foreshore/tidal Shoreface Foreshore/tidal Shoreface Shoreface O¡shore Shoreface O¡shore O¡shore Shoreface O¡shore
s 25 s 15 s 15 s 15 s 30 s 15 s 25 ? s 25 s 15 s 25 s 25 s 15 s 30 s 30 s 30 s 30 s 25 s 30
Boreal Boreal Ubiquitous Boreal Arctic Arctic Arctic Arctic Arctic Arctic
Foreshore/tidal Foreshore/tidal Foreshore/tidal Shoreface Shoreface Shoreface Shoreface Foreshore/tidal Shoreface Foreshore/tidal
Subarctic Boreal Arctic Boreal Boreal Boreal Boreal Boreal Boreal Boreal Lusitanic Boreal Boreal
Shoreface Foreshore/tidal Foreshore/tidal Shoreface Foreshore/tidal Foreshore/tidal O¡shore Shoreface Foreshore/tidal Foreshore/tidal Foreshore/tidal O¡shore Foreshore/tidal
zone zone zone
zone zone
zone zone zone
zone zone
zone zone zone zone
zone zone zone zone
PALAEO 2866 25-7-02 Cyaan Magenta Geel Zwart
s 15 s 15 s 25 s 25 s 25 s 25 s 25 s 25 s 25 s 30 s 15 s 15 s 30 s 15 s 15 s 15 s 30 s 30 s 15 s 15 ? s 15 s 30
Arkhangelsk region
s 15 s 15
Watershed area
O¡shore Shoreface
Petrozavodsk area
Boreal Ubiquitous
St. Petersburg area
Depth
Vistula
Biogeographical characteristics
Belt Sea
Temperature index
Wadden Sea
Species
Salinity tolerance (x)
Table 1(Continued).
F F
F F F
F F
F F
F
F F F F F F
F
F
F
F F
F F F F F F
F
F
F
F
F
F
F F
F F F F
F F F F F
F
F
F
F
F F F
F F
?
F F F
F
F
F
280
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Shoreface ? Foreshore/tidal Foreshore/tidal Foreshore/tidal Foreshore/tidal Foreshore/tidal Foreshore/tidal Foreshore/tidal Shoreface Foreshore/tidal Foreshore/tidal
s 15 ? s 15 s 25 s5 s 15 s 30 ? s 15 s 15 s 15 s 15
F
F
F
F
F
F
F
61.0 68.0 61.0 74.0 100.0 99.6
Boreal Boreal Boreal Subarctic Ubiquitous Arctic
Shoreface Shoreface O¡shore O¡shore O¡shore Changing depth requirements Changing depth requirements Shoreface Changing depth requirements O¡shore Changing depth requirements Shoreface Foreshore/tidal zone O¡shore Foreshore/tidal zone Foreshore/tidal zone Foreshore/tidal zone Foreshore/tidal zone Changing depth requirements Foreshore/tidal zone Foreshore/tidal zone Shoreface Foreshore/tidal zone O¡shore Shoreface O¡shore O¡shore Shoreface Shoreface
s 15 s 30 s 15 s 25 s 15 s 15
F
F
Nuculana minuta (Mu«ller, 1776)
80.0
Yoldia hyperborea Torell, 1859 Yoldiella frigida (Torell, 1859)
99.7 100.0
Yoldiella intermedia (Sars, 1865) Yoldiella lenticula (MUller, 1842)
99.7 99.7
Subarctic Arctic Ubiquitous Arctic Arctic
Portlandia arctica (Gray, 1824) 99.7 Portlandia aestuariorum Mossevitch, 1928 99.7 Bathyarca glacialis (Gray, 1824) 99.7 Glycymeris glycymeris (Linne¤, 1758) 62.5 Mytilus edulis Linne, 1758 75.3 Mytilaster lineatus (Gmelin, 1791) 40.0 Mytilaster cf. minimus (Poli, 1795) 48.0 Crenella decussata (Montagu, 1808) 85.1
Arctic Arctic Arctic Boreal Subarctic Lusitanic Lusitanic Subarctic
Musculus discors (Linne¤, 1767) Musculus tumida (Hanley, 1843) Modiolus modiolus (Linne, 1758) Modiolula phaseolina (Philippi, 1844) Pecten maximus (Linne¤, 1758) Aequipecten opercularis (Linne¤, 1758) Arctinula greenlandica (Sowerby, 1842) Chlamys islandica (Mu«ller, 1776) Chlamys varia (Linne¤, 1758) Heteranomia squamula (Linne¤, 1758)
Ubiquitous Boreal Boreal Boreal Boreal Boreal Arctic Subarctic Boreal Subarctic
100.0 70.5 74.9 71.3 67.0 69.0 99.7 85.1 64.0 78.2
zone zone zone zone zone zone zone zone zone
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Arkhangelsk region
Boreal Lusitanic Boreal Boreal Subarctic Boreal Boreal Lusitanic Boreal Ubiquitous Boreal Boreal
Watershed area
Vistula
65.0 40.0 70.0 73.3 80.0 70.0 70.5 53.0 67.0 100.0 68.0 71.3
Petrozavodsk area
Depth
St. Petersburg area
Biogeographical characteristics
Belt Sea
GASTROPODA Ebala nitidissima (Montagu, 1803) Ebala pointeli (de Folin, 1868) Acteon tornatilis (Linne¤, 1758) Diaphana minuta Brown, 1827 Retusa obtusa (Montagu, 1803) Retusa truncatula (Bruguie're, 1792) Retusa umbilicata (Montagu, 1803) Haminoea navicula (da Costa, 1778) Philine aperta (Linne¤, 1767) Cylichna alba (Brown, 1827) Cylichna cylindraceae (Pennant, 1777) Akera bullata (Mu«ller, 1776) BIVALVIA Nucula nitidosa Winckworth, 1930 Nucula nucleus (Linne¤, 1758) Nucula sulcata Bronn, 1831 Nuculoma corticata (MUller, 1842) Nuculoma tenuis (Montagu, 1808) Nuculana pernula Mu«ller, 1779
Temperature index
Wadden Sea
Species
Salinity tolerance (x)
Table 1(Continued).
F F
? F F
F
F
F
F
F
F
F
F
?
F F
F F F F
F
s 15
F
s 15 s 30
F
s 30 s 30
F
s 25 65 s 15 s 30 65 ? ? s 15 s 15 s 15 s 15 s 30 s 15 s 15 s 15 s 25 s 30 s 15
F
F
F
F
F
F
F
F F F
F
F
?
F
F
F
F
F
F F F
F F F
F
F
F
F
F F F F
F
F
F
F F
F
F
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281
Astarte crebricostata Mc Andrews and Forbes, 1847 Astarte crenata (Gray, 1824) Astarte elliptica (Brown, 1827) Astarte montagui montagui Dillwyn, 1817 Astarte montagui striata (Leach, 1819) Astarte sulcata (da Costa, 1778) Acanthocardia echinata (Linne, 1758) Acanthocardia paucicostata (Sowerby, 1841) Parvicardium elegantulum (MUller, 1842) Parvicardium exiguum (Gmelin, 1791) Parvicardium ovale (Sowerby, 1840) Parvicardium scabrum (Philippi, 1844) Plagiocardium papillosum (Poli, 1795) Laevicardium crassum (Gmelin, 1791) Cerastoderma edule (Linne, 1758) Cerastoderma glaucum (Poiret, 1789) Ciliatocardium ciliatum (Fabricius, 1780) Serripes groenlandicus (Bruguie're, 1789) Mactra stultorum (Linne¤, 1758) Spisula elliptica (Brown, 1827) Spisula solida (Linne¤, 1758) Spisula subtruncata (da Costa, 1778) Lutraria lutraria (Linne¤, 1758) Ensis ensis (Linne¤, 1758) Ensis siliqua (Linne¤, 1758) Phaxas pellucidus (Pennant, 1777) Tellina donacina Linne¤, 1758 Tellina fabula Gmelin, 1791 Tellina tenuis da Costa, 1778 Macoma balthica (Linne, 1758) Macoma calcarea (Gmelin, 1791)
Arctic
Shoreface Shoreface Foreshore/tidal zone Shoreface O¡shore Shoreface O¡shore O¡shore O¡shore Foreshore/tidal zone Foreshore/tidal zone Changing depth requirements Shoreface
?
99.6 100.0 85.1 81.0 72.3 72.9 50.0
Arctic Ubiquitous Subarctic Subarctic Boreal Boreal Lusitanic
O¡shore Shoreface Shoreface Shoreface O¡shore Shoreface ?
? s 15 s 15 ? s 30 s 15 ?
73.9 70.0 73.1 70.5 56.0 62.0 71.0 71.0 99.7 99.7 58.0 73.1 57.0 67.5 56.0 62.0 63.5 70.0 55.0 68.5 70.0 85.1 100.0
Boreal Boreal Boreal Boreal Boreal Boreal Boreal Boreal Arctic Arctic Boreal Boreal Boreal Boreal Boreal Boreal Boreal Boreal Boreal Boreal Boreal Subarctic Ubiquitous
O¡shore Shoreface Shoreface Shoreface Shoreface Shoreface Foreshore/tidal zone Foreshore/tidal zone Shoreface Shoreface Foreshore/tidal zone Shoreface Shoreface Foreshore/tidal zone Foreshore/tidal zone Foreshore/tidal zone Foreshore/tidal zone Shoreface O¡shore Foreshore/tidal zone Foreshore/tidal zone Foreshore/tidal zone Changing depth requirements
? s 15 s 15 s 30 ? s 30 s5 65 s 15 s 15 s 30 s 15 s 30 s 15 s 30 s 25 s 25 s 15 s 15 s 30 s 15 65 s5
99.7
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s 15 s 25 s 25 ? s 25 s 25 s 15 s 30 s 25 s 15 s 30 s5
F
Arkhangelsk region
F
Watershed area
Boreal Boreal Lusitanic Lusitanic Boreal Arctic Boreal Boreal Boreal Boreal Boreal Arctic
Petrozavodsk area
70.5 65.5 53.0 51.0 71.3 99.7 62.0 60.5 70.0 70.0 72.0 99.6
Depth
St. Petersburg area
BIVALVIA Pododesmus patelliformis (Linne¤, 1761) Ostrea edulis Linne¤ 1758 Loripes lucinalis (Lamarck, 1818) Lucinella divaricata (Linne¤, 1758) Lucinoma borealis (Linne¤, 1767) Axinopsida orbiculata (Sars, 1878) Thyasira £exuosa (Montagu, 1803) Hemilepton nitidum (Turton, 1822) Montacuta ferruginosa (Montagu, 1808) Mysella bidentata (Montagu, 1803) Turtonia minuta (Fabricius, 1780) Astarte borealis (Schumacher, 1817)
Vistula
Biogeographical characteristics
Belt Sea
Temperature index
Wadden Sea
Species
Salinity tolerance (x)
Table 1(Continued).
F
F
F
F
F
F
F F F
F
F
F
F
F
F
F
F
F
F F F
F
F
F
F F
F
F
F
F
F F F
F
F
?
F
F
F
F
F
F F
F
F
F
F F F F
F
F F
F
F
F
F
F
F
F F
F F F
F
? F
F
F
F
F F
F
F
F F F F F
F
F
F
F
F F
F
F
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Balanus improvisus Darwin, 1854 Semibalanus balanoides (Linne¤, 1758)
69.0 80.4
Foreshore/tidal zone Shoreface Shoreface Changing depth requirements Foreshore/tidal zone Foreshore/tidal zone
s 30 s5 s5 s 25
F
65 s 15
F
Boreal Subarctic
with information on their requirements in relation to temperature, salinity and depth, as described below. With few exceptions the faunas come from shoreface habitats and re£ect surface temperature and salinity. The taxonomical nomenclature generally follows Le Renard (1997) and
F
F
F
F
F
F
F
Arkhangelsk region
Boreal Ubiquitous Subarctic Subarctic
F
Watershed area
s 15 s 30 s5 s5 s 30 s 25 65 s5 s 30 s 25 s 25 s 30 s 25 s 30 s 30 ? s 15 s 15 s5 s 15 s5 s 25 s 15 s 15 s5 s 25 s 15 s 25 s 25 s 15
Petrozavodsk area
Shoreface Shoreface Foreshore/tidal zone Shoreface O¡shore O¡shore Shoreface Foreshore/tidal zone O¡shore Shoreface O¡shore Shoreface O¡shore Shoreface Shoreface ? Shoreface O¡shore Shoreface O¡shore Shoreface Shoreface O¡shore Shoreface Shoreface O¡shore Shoreface O¡shore Foreshore/tidal zone Shoreface
St. Petersburg area
Lusitanic Boreal Boreal Boreal Boreal Boreal Lusitanic Boreal Boreal Boreal Boreal Boreal Boreal Boreal Boreal ? Boreal Boreal Ubiquitous Boreal Ubiquitous Boreal Boreal Boreal Boreal Boreal Arctic Boreal Boreal Boreal
Vistula
Depth
Belt Sea
BIVALVIA Gastrana fragilis (Linne¤, 1758) 48.0 Donax vittatus (da Costa, 1778) 58.5 Scrobicularia plana (da Costa, 1778) 62.0 Abra alba (Wood, 1802) 67.5 Abra nitida (Mu«ller, 1776) 70.5 Abra prismatica (Montagu, 1803) 70.5 Abra segmentum (Re¤cluz, 1843) 47.0 Arctica islandica (Linne, 1767) 72.4 Kelliella abyssicola (Forbes, 1844) 69.0 Chamelea gallina (Linne¤, 1757) 71.3 Timoclea ovata (Pennant, 1777) 71.3 Gouldia minima (Montagu, 1803) 60.0 Dosinia lupinus (Linne¤, 1758) 70.0 Tapes decussata (Linne¤, 1758) 62.0 Paphia aurea (Gmelin, 1791) 63.5 Paphia senescens (Cocconi, 1873) 0.0 Venerupis senegalensis (Montagu, 1803) 70.0 Mysia undata (Pennant, 1777) 67.5 Mya truncata Linne¤, 1758 100.0 Corbula gibba (Olivi, 1792) 70.5 Hiatella arctica (Linne¤, 1767) 100.0 Panomya arctica (Spengler, 1793) 73.3 Saxicavella je¡reysi Winckworth, 1930 68.0 Barnea candida (Linne¤, 1758) 64.0 Zirfaea crispata (Linne, 1767) 71.3 Thracia convexa (Wood, 1815) 60.5 Thracia myopsis (MUller, 1842) 99.7 Thracia papyracea (Poli, 1791) 68.0 Thracia villosiuscula (Macgillivray, 1827) 60.0 Verruca stroemia (Mu«ller, 1776) 73.3 CIRRIPEDIA Chtamalus stellatus (Poli, 1791) 55.0 Balanus balanus (Linne¤, 1758) 100.0 Balanus crenatus (Bruguie're, 1789) 75.3 Balanus hameri (Ascanius, 1767) 85.1
Biogeographical characteristics
Wadden Sea
Temperature index
Species
Salinity tolerance (x)
Table 1 (Continued).
F
F
F
F F
F
F
F
F
F
F
F
F
F F
F
F
F
F
F
F
F
F
F F F
F
F
F F
F F
F
F
F
F
F
F F
F F
F
F
F
F
F
F
F F
F F F F
F
F
F F
F F
F
F
F
F F
Poppe and Goto (1991, 1993), who give comprehensive synonyms. 1.4. Biogeographical zones The species are classi¢ed according to their
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283
Fig. 2. Distribution of marine biogeographical zones in the North Atlantic. (A) Present, (B) Early Eemian.
northern penetration along the European coasts (Fig. 2). The biogeographical terms are here used in an oceanographic sense, based on water mass characteristics, and deviate slightly from Feyling-Hanssen’s (1955) often-used scheme, which was based on the occurrence of indicator species along the Norwegian coast. The Lusitanian zone is characterised by dominance of warm Atlantic water as seen especially in the high winter sea surface temperatures, s 9‡C, and summer temperatures higher than 16‡. The boundary for this zone, which marks the northernmost occurrence of a number of Mediterranean species, is at the southwestern entrance to the English Channel (Ekman, 1953). A notable representative of this fauna is Lucinella divaricata, which penetrated far into the Baltic during the Eemian. Also the Boreal zone is strongly in£uenced by warm Atlantic surface water. Summer sea surface temperatures are higher than 8‡C and the area is year-round ice-free. This zone is bounded to the north and east by the limit for regular occurrence of winter sea ice at the Murman coast. The zone extends for almost 2000 km from S to N along the North European coast, and is characterised throughout by its uniform shallow water faunas with Arctica islandica, Spisula elliptica, Cerastoderma edule, and Zirfaea crispata.
During the Eemian this zone extended through the Barents and Kara Seas to Taymyr (Fig. 2). To the north and east of this, the subarctic zone is the zone of mixing of Polar and Atlantic water. Summer sea surface temperatures are higher than 3‡C and although winter sea ice is present, there is at least a 2-month ice-free period. The foreshore and intertidal fauna (e.g. Mytilus edulis, Semibalanus balanoides, Macoma balthica) has its northern limit in this area. Further to the north and east cold polar water and long lasting ice cover characterise the arctic zone. The long lasting ice cover makes stationary life on the foreshore di⁄cult and the foreshore fauna is missing. The upper shoreface is inhabited by such species as Portlandia arctica, Arctinula greenlandica, and Bathyarca glacialis. The cold polar water from the Arctic basin sinks down and occurs as deep water far to the south. The arctic species may therefore occur at greater depths even in the boreal zone. Finally, some species have a wide tolerance towards temperature and are ubiquitous in the entire region. 1.5. Temperature As an expression of temperature demands the lusitanian, boreal, and subarctic species are in-
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dexed after their northern boundaries (Table 1). Along the south^north trending European coastline this is expressed as latitude (72.6 = 72‡60PN). Along the west^east trending Russian arctic coastline it has been adjusted by calibration with species with a northern limit on Svalbard. The arctic species are indexed according to their southern boundaries (90.72Wsouthern limit at 72‡N). The distribution and temperature demands are based on information from Jensen and Spa«rck (1934), Stephensen (1933, 1938), Gayevskaja (1948), Filatova (1957), Tarasov and Zevina (1957), Zenkevitch (1963), Tebble (1966), Scarlato (1981, 1987), HUisZter (1986), Graham (1988), Seaward (1990, 1993), Golikov (1995), and Bratlegaard and Holthe (1997). 1.6. Salinity Life in water with reduced salinity is di⁄cult for most marine organisms, and the lower the salinity the lower the number of species that can endure it. Also, reduced in£ow of oceanic water means stronger in£uence of atmospheric temperatures and larger annual temperature variations, making life di⁄cult for both warmth and cold demanding species. In Table 1 the species are grouped after their tolerance to reduced salinity into euhaline ( s 30x), and polyhaline ( s 25x), b-mesohaline ( s 15x), a-mesohaline ( s 5x), and oligohaline ( 6 5x). The species’ salinity demands are based on information from Scarlato (1981, 1987) and Golikov (1995), and their occurrence on the salinity gradient from the open ocean through the North Sea and into the Baltic (Seaward, 1990, 1993; Jensen and Knudsen, 1995; KUie et al., 2000). Common brackish-tolerant species are Mytilus edulis, Macoma balthica, and Balanus improvisus. In the Arctic Portlandia aestuariorum inhabits, as the only mollusc, estuaries with salinities down to less than 1x (Scarlato, 1981; Petryashov et al., 1999). Most low salinity-tolerant species can thrive in oceanic water and estimating salinity depends on the species with the least tolerance to brackish conditions. Among the most demanding oceanic representatives can be mentioned Mactra stultorum and Tapes decussatus.
1.7. Depth The benthic species are not dependent on water depth. However, since such environmental parameters as wave action, temperature variation, and photosynthesis are related to depth this is a major determining factor for their distribution. In the shallow water wave action and tide are probably the most important restrictive factors, and the species are here categorised according to their shallowest depth on the shore plane from foreshore/tidal to shoreface and o¡shore. Many arctic molluscs prefer to live high on the shoreface in the Arctic, but extend their distribution to o¡shore environments in the boreal zone, and therefore have changing depth requirements. The depth parameter is used here to indicate habitat diversity in each area. 1.8. Age We date and correlate the Eemian marine sediments by using published pollen diagrams from marine sediments, supplemented with a new diagram from Petrozavodsk. The Eemian in northern Europe is characterised by a uniform development of vegetation, and similar pollen zones can be identi¢ed in the entire area of deciduous forest (Zagwijn, 1996), which extended from the Atlantic coasts eastwards to Moscow and to Lake Onega in the north (Grichuk, 1984). In the Early Eemian the forest trees immigrated very rapidly and seem to have arrived everywhere in the same order of succession. Since the whole immigration period was short ( 6 3 ka), the di¡erence in arrival time of a species to di¡erent parts of the region may not have amounted to more than a few centuries (Zagwijn, 1996; Cheddadi et al., 1998). After the immigration rush came the period of stability, beginning with the Carpinus zone. In this period the di¡erences in timing of pollen zones are probably much greater, and it would be unlikely if the period of deciduous forest in the north of Russia was not considerably shorter than in the south, whereas the subsequent taiga stages, the Picea and Pinus zones, were longer. Since we do not know the migration routes for the trees, this error cannot be estimated, but it
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may not a¡ect our results seriously, because the events discussed here mainly took place in the Early Eemian. The durations of the pollen zones in Table 2 are Mu«ller’s (1974) estimates, derived from the counting of annual layers in lake sediments from North Germany. The duration of the early pollen zones, the ¢rst 2.1 ka, was done with high precision, but the duration of the remaining part of the Eemian had to be made by extrapolation and is much more uncertain (Mu«ller, 1974). Tentatively we suggest absolute ages for the pollen zones (Table 2). This is based on a correlation of Zagwijn’s (1983) pollen-dated sea level curve from the southern North Sea with curves for eustatic sea level change based on U^Th dating of corals (Stirling et al., 1998; Esat et al., 1999; McCulloch et al., 1999; Henderson and Slowey, 2000). The correlation ¢x-point in the two records is the attainment of the Eemian sea level highstand, here estimated as ca. 500 yr after the beginning of the Carpinus zone and dated on coral reefs to 128 [ 1 ka (McCulloch and Esat, 2000; Shackleton, 2000). From this we arrive at tentative ages for the Eemian pollen zones in northern Europe (Table 2)1 . This estimate is tentative and ignores isostatic e¡ects, which could have terminated the transgression in the area before the attainment of the global highstand, and shifted the beginning of the ‘pollen-Eemian’ further back in time if isostatic uplift was faster than sea level rise; or the transgression could have lasted longer if the area was subjected to forebulge collapse. Disregarding these factors may to some extent be justi¢ed by sea level curves (Esat et al., 1999; McCulloch et al., 1999; Israelson and Wohlfahrt, 1999), which indicate that sea level rise was so rapid that it is likely to have overridden the isostatic e¡ects, and the change from very high to zero rate of sea level rise was
1
This estimate has the consequence that the ‘pollen-Eemian’ here ends at 120.5 ka, and not at the well established age of ca. 115 ka. It is beyond the scope of this paper to discuss the cause for this discrepancy. However, it should be noted that the duration of the Late Eemian pollen zones is based on extrapolation, which could have minimised their durations (Mu«ller, 1974).
285
abrupt, suggesting that maximum transgression was coeval with the attainment of the global highstand. This is supported by the fact that regression in many areas within the region began in the Carpinus zone.
2. Description of areas Table 1 gives lists of molluscs and cirripeds from each area along the seaway shown on Fig. 1. Notes are given below of their type of occurrence, pollen analytical results, and outline of marine inundation. The duration of marine inundation at each site is shown in Table 2. In some areas, notably the Gulf of Bothnia, there are no mollusc records, and the marine environment is deduced from diatoms. 2.1. The Wadden Sea This area comprises the southern North Sea, southwestern Jutland and northwestern Germany. Eemian marine sediments are known from numerous borings along the 130 km coast, which lies outside the margin of Weichselian glaciation. The sedimentary successions comprise basal sandy shoreface facies, ‘senescens sand’, named after Paphia senescens, overlying limnic sediments, and overlaid by thick ‘Turritella clay’, named after Turritella communis, or ‘Olander beds’, which are strati¢ed ¢ne sand and clay, probably tidal (Dittmer, 1951; Temmler, 1995). Further inland the sediments show great variability and re£ect a wide variety in hydrographic conditions in this area of narrow straits and broads (Gripp, 1964; Kosack and Lange, 1985; Temmler, 1995). In general the successions show a transition from limnic to brackish to marine to brackish to limnic, and re£ect a complete transgressive^regressive cycle. The mollusc fauna in Table 1 represents all these sedimentary facies and is based on the studies of Heck and Brockmann (1950), Dittmer (1951, 1954), Jaeckel (1963), Hinsch (1985), and especially Nordmann (1928). The rich fauna owes partly to the diversity in habitats and hydrography, but also to the high salinity.
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Table 2 Correlation and duration of pollen zones along the White Sea^Baltic seaway
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PALAEO 2866 25-7-02 Cyaan Magenta Geel Zwart Dark shading marks maximum duration of White Sea^Baltic seaway. (1) Based on the listed durations and correlation between pollen- and U^Th-dated sea level curves (Zagwijn, 1983; Esat et al., 1999; Henderson and Slowey, 2000).
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In Schleswig-Holstein the transgression came in the Betula^Pinus zone (Menke, 1985), as evidenced by drowning surfaces and transgression conglomerates with Ostrea edulis showing that both sea surface temperatures and salinity were high at the very beginning of the Eemian (Hinsch, 1985; Knudsen, 1985; Meijer and Preece, 1995). This implies the English Channel was open at the beginning of the Eemian and transported more Atlantic water into the North Sea than during the Holocene (Meijer and Preece, 1995). An even warmer and more saline fauna is known from Holland, but not pollen analytically dated within the Eemian (Spaink, 1958; Meijer and Preece, 1995). The large proportion of oceanic species, which do not appear in the North Sea at present (e.g. Cerithiopsis tubercularis, Odostomia conoidea, Tapes decussatus), show that maximum salinity, s 30x, was reached in the Corylus, Taxus/Tilia, and early Carpinus zone. After this, regression set in and during the Picea zone the sea retreated from this area. The Eemian sea inundated valleys and depressions and gave the coastline a complex outline with small irregular inlets penetrating deep into the country and land areas extending out into what is now the southern North Sea, maybe as far as Helgoland (Dittmer, 1951, 1954; Gripp, 1964; Behre, 1970; Konradi, 1976; Kubisch and Scho«nfeld, 1985; Knudsen, 1994; Friborg, 1996). In the area of the Kiel Canal in situ marine sediments outline a winding sound with broads and narrows extending across Jutland from the Friesian islands eastwards to areas behind the Weichselian ice margin, less than 20 km from the present Baltic coast. This made Kosack and Lange (1985) suggest the existence of a seaway from the North Sea to the Baltic. 2.2. Kattegat and the Belt Sea The inner Danish coasts and those of northern Germany as far as Mecklenburg^Vorpommern were overridden by ice during the Weichselian, and almost all occurrences of marine Eemian sediments are dislocated clasts. The fauna list in Table 1 is compiled from Nordmann (1928),
287
Heck and Brockmann (1950), Gehl (1961), Madsen (1965), and Houmark-Nielsen (1994). The Cyprina clay, named after Cyprina (now Arctica) islandica, is exposed in coastal cli¡s around the western Baltic in Denmark and northern Germany, possibly as far east as Ru«gen and Usedom (Konradi, 1976; Kubisch and Scho«nfeld, 1985; Ho«£e et al., 1985; Mu«ller et al., 1995; Ru«hberg et al., 1995). The lower contact is usually a drowning surface overlying limnic sand or clay. At its top the Cyprina clay is usually truncated by glacial or glaci£uvial sediments, only in the cli¡s at Gammelmarke does the clay grade into shoreface sand rich in shells, the ‘Tapes sand’, named after Tapes (now Paphia) senescens with a fauna similar to that in the senescens sand in Schleswig-Holstein. Dislocated clasts containing a deep-water fauna similar to the Turritella clay of Schleswig-Holstein are also known from a few sites (\dum, 1933; Houmark-Nielsen, 1994). The faunas contain oceanic species, which nowadays are absent from the North Sea and Kattegat (Turbonilla lactea, Gouldia minima, Hemilepton nitidum) or they live in the North Sea and northern Kattegat, but not in southern Kattegat and the Belt Sea (Caecum glabrum, Mactra stultorum). All these oceanic species are, however, known only from the Gammelmarke cli¡s in the westernmost Baltic (Madsen et al., 1908), but in the eastern Belt Sea polyhaline species such as Ostrea edulis, Turritella communis, and Aporrhais pespelicani show that salinities here also were higher than those attained during the Holocene and the common occurrence of Lucinella divaricata shows that temperatures were higher. This shows that the Belt Sea, after a brief brackish phase, had salinities of 25^30x. At the western edge of the basin the salinity was at least periodically even higher. At Ristinge the marine inundation has recently been dated to the beginning of the Quercus zone, and the marine inundation lasted ca. 3.4 ka, until the early Carpinus zone (Kristensen et al., 2000). Ostracodes, foraminifera and dino£agellates indicate an initial brackish phase followed shortly afterwards by more saline conditions, which culminated at the start of the Carpinus zone
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(Table 2), although fully marine conditions were never attained (Head and Gibbard, 2000). Also in eastern Schleswig-Holstein and Mecklenburg the marine inundation took place before the Carpinus zone and the retreat came in the Picea zone (Heck and Brockmann, 1950; Gehl, 1961).
higher than 25x (Fig. 3). By the end of the Carpinus zone salinity dropped, as evidenced by the low diverse Mytilus edulis^Cerastoderma glaucum assemblages.
2.3. Western Baltic
In the Neva lowland and areas around Lake Ladoga Eemian marine sediments consist of dark organic and gas rich laminated clay and silt with thin layers of ¢ne sand, known both from numerous borings and from temporary exposures in clay pits (MalakhovskiY et al., 1989; Krasnov et al., 1995). The sediments occur both in situ and as glacially dislocated clasts. They represent o¡shore or lagoonal mud deposited as basin ¢ll in drowned valleys and depressions (Znamenskaia and Cherminisova, 1962; Malakhovskiy et al., 1989). The best-known occurrence is at Mga. Pollen diagrams from this and nearby Rybatskoe show that the marine drowning began early in the Eemian, in the Pinus^Ulmus zone, and the marine phase lasted until the Pinus zone (Table 2 and Znamenskaja, 1959; Lavrova, 1962). The mollusc faunas from Mga and other sites have been described by Skorokhod (1932) and Lavrova (1961, 1962), and are preserved in the Geological Museum in St. Petersburg where we have had the opportunity to study them. With a total of only eight species the faunas are notable for their low diversity and lack of warm species. They are characterised especially by Portlandia arctica and Portlandia aestuariorum (Skorokhod, 1932). The fauna shows increasing and then decreasing salinity upwards in the sediments with maximum salinity being achieved prior to the Corylus zone (Fig. 3). This is shown by the transition from predominance of P. arctica in the low-
During the Eemian the Vistula valley was inundated by the sea up to 70 km from its present mouth in the Danzig Bay. The sediments, considered to be in situ, are known from rare exposures and many borings and comprise shoreface sand and o¡shore clay (Makowska, 1986, 1991). The mollusc faunas in Table 1 have been compiled from a number of sites (Brodniewics, 1960, 1969; Makowska, 1986), especially Elblag and Nowiny, which display normal transgressive^regressive sequences. The domination of Corbula gibba throughout indicates a protected fjord environment. In the Nowiny boring the marine episode began at the onset of the Quercus zone and lasted until some time into the Picea zone (Makowska, 1986). Paphia senescens was among the very ¢rst immigrants, and Lucinella divaricata followed immediately after, showing that the connection to the North Sea was established and sea surface temperatures already were higher than Holocene. Retusa umbilicata and Abra nitida are sparsely present throughout, and indicate that salinity culminated in the Quercus zone, periodically rising to over 30x. The poor development of Ostrea and Venerupis senegalensis (earlier pullastra, Brodniewics, 1960, 1969), on the other hand, shows that conditions were not optimal for them, suggesting that salinity probably generally was not
2.4. Eastern Baltic
Fig. 3. The Baltic^White Sea region at the peak of marine inundation in the Early Eemian, tentatively dated to the period 132^ 130 ka. Red arrows mark entranceways discussed in the text. The development of salinity at each locality is shown in columns at the base of the ¢gure. The columns represent the duration of the Eemian divided into pollen zones. The outline of the seaway is based on an assessment of maps in: Apukhtin and Krasnov, 1966; Atlas Karelskoi ASSR, 1989; Biske, 1959; Biske and Devyatova, 1965; Devyatova, 1961, 1982; Devyatova and Loseva, 1964; Forsstro«m et al., 1988; Friborg, 1996; Gehl, 1961; Goretskyi, 1949; Gripp, 1964; Ho«£e et al., 1985; International Quaternary map of Europe, 1970^1971; Knudsen, 1994; Konradi, 1976; Kosack and Lange, 1985; Lavrova, 1960, 1961; Legkova and Schukin, 1967; Lundqvist and Robertsson, 1994; Makowska, 1986; Malakhovskiy et al., 1989; Ru«hberg et al., 1995; Seidenkrantz et al., 2000; Temmler, 1995; Znamenskaia and Cherminisova, 1962. Bathymetry is modelled from present depths and Seidenkrantz and Knudsen (1997).
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290
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Fig. 4. Smear of Portlandia aestuariorum in laminated clayey silt from Mga (from collections in the Geological Museum, St. Petersburg).
er part to predominance of the more brackish P. aestuariorum from the beginning of the Carpinus zone (Skorokhod, 1932). Portlandia is present from the earliest phase (Lavrova, 1962) and is concentrated in smears and thin lenses, probably indicating mass death and gentle in-habitat reworking (Fig. 4). The other components of the fauna occur as single individuals scattered in the sediment. They are present only at the salinity maximum in the middle part of the sequence and comprise Macoma calcarea, thin-shelled small Mytilus edulis and a few Cerastoderma, originally identi¢ed as C. edule, but their triangular appearance makes it more likely that they belonged to the more brackish Cerastoderma glaucum. These shells, together with leaf fragments of Zostera marina, were probably redeposited from the nearby foreshore (Lavrova, 1962). M. calcarea, C. glaucum and a single worn Littorina littorea, as well as the small and thin M. edulis suggest surface salinity at ca. 15x when it peaked in the middle sequence. Portlandia arctica, on the other hand, generally requires salinity of more than 25x (e.g. Mossevitch, 1928; Scarlato, 1987), suggesting strati¢cation between low saline surface water and denser bottom water. We suggest that the high-saline and warm planktonic diatoms (Brander, 1937; Znamenskaia and Cherminisova, 1962) could represent brief injections of warm saline water surface water from the Baltic, such as those that control the salinity of the Baltic at present (Bo«rngen et al., 1990). The scarcity of molluscs, their concentration in
smears or in sand laminae, and the high organic content of the sediments indicate that anoxic bottom conditions were frequent, but interrupted by short intervals of oxygenation. Anoxic conditions would also explain the rarity of benthic and attached diatoms, which induced Znamenskaia and Cherminisova (1962) to assume a water depth of 80^100 m over the sites. At Prangli, at the mouth of the Gulf of Finland, an Eemian pollen diagram from marine sediments from a boring shows a gradual change from Late Saalian limnic and brackish phase and into ‘normal marine’ conditions in the Early Eemian (Liivrand, 1991). A sudden change from brackish to marine took place by the end of the Pinus^Ulmus zone. At this place in the core mollusc shells also appear to have their ¢rst appearance, which we consider to mark the marine inundation. The marine conditions continue into the early Picea zone, and are ended by a hiatus (Fig. 3). These results have recently been con¢rmed and expanded by pollen^foraminifer- and diatom analyses of Eemian marine sediments in three borings in coastal Latvia and the Riga Bay (Kalnina, 2001). In the boring at Plasumi the marine sediments cover the entire Eemian, beginning and ending with cold brackish water facies in the Late Saalian and Early Weichselian. Diatoms and foraminifera indicate that water depth and salinity, s 18x based on diatoms, culminated before the Carpinus zone, when regression set in (Kalnina, 2001). On the eastern shore of Lake Ladoga marine
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reworked megaclasts of Eemian marine sediments have been encountered in borings and pollen diagrams give minimum ages for the duration of the marine inundation of this basin, lasting from the Quercus and into the early Picea zone (Devyatova, 1982). 2.5. The Gulf of Bothnia In situ marine sediments in borings at three localities in Ostrobothnia were deposited immediately after deglaciation or overlie lacustrine sediments during fall in relative sea level (Eriksson et al., 1980, 1999). Pollen diagrams and correlation with sea level history indicate that the marine inundation began after the immigration of Quercus and lasted at least until within the Picea zone. Initially, high percentages of polyhalobous diatoms indicate salinities higher than 17x. Upwards in the sediments a decline in salinity began before the onset of the Carpinus zone (Fig. 3). In a boring at Norra Sanna«s in central Sweden marine Eemian clay is topped by clay with fresh water diatoms, and the marine inundation lasted from the Quercus zone until some time before the beginning of the Carpinus zone (Robertsson et al., 1997). Reworked brackish-marine sediments occur also at other sites and indicate that salinity in the Early Eemian was higher than Holocene and sea level fall began before the immigration of Picea (Robertsson and Garc|¤a Ambrosiana, 1992; Robertsson, 2000). 2.6. Karelia, Petrozavodsk Marine Eemian sediments are known from borings in isolated sediment pockets preserved in depressions of the Precambrian basement. In the city of Petrozavodsk borings have shown marine Eemian sediments considered to be in situ at 40 m above sea level (asl). They show continuous development from glaciolacustrine clay to marine sediments and lacustrine sand and till (Wollosovich, 1908; Lukashov, 1982). The lower clay contains plant remains and is dominated by Macoma calcarea, while the upper sandier facies is bioturbated and dominated by Portlandia aestuariorum and thin-shelled Mytilus edulis. The mollusc fauna
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suggests salinities at 10^15x, based on M. calcarea, at the base to somewhat lower at the top. The bioturbation shows that anoxia was not prevalent as in the St. Petersburg area (Wollosovich, 1908; Skorokhod, 1932). Pollen diagrams from two borings in the city of Petrozavodsk contain su⁄cient pollen of deciduous trees for the record to be correlated with the West European pollen stratigraphy (Fig. 5 and Lukashov, 1982). These results show that the marine inundation took place in the Early Eemian, in the Pinus^Ulmus zone. In the diagram presented here the marine episode lasted until the end of the Corylus zone (Fig. 5), while the earlier diagram by Devyatova (in Lukashov, 1982) sets the boundary somewhat later in the upper part of the following Taxus^Tilia zone. By palynological correlation this corresponds to a duration of ca. 2.5 ka, tentatively from ca. 132.4 to ca. 130 ka. Petrozavodsk is close to the continental watershed towards the White Sea, and the sedimentary sequence can be considered as a monitor for marine over£ow from the White Sea, implying that the over£ow and the seaway existed only in the Early Eemian. After this the connection was severed to the north of Lake Onega, but the Ladoga^ Onega Lake basin probably continued as a brackish arm of the Baltic for some time. The outline of the Eemian basin in Fig. 3 is reconstructed using evidence from Apukhtin and Krasnov (1966), Atlas Karelskoi ASSR (1989), and Devyatova (1961, 1982). The two lake basins were connected by a wide shallow sound, 50 km across at the narrowest, and with scattered islands. In its northern end the Onega basin opened up towards the White Sea. 2.7. The continental watershed area Between Lake Onega and the White Sea shore 150 km to the north. The marine Eemian sediments occur as isolated patches ¢lling up depressions in the bedrock surface (Zemlyakov, 1936; Goretskyi, 1949; Biske, 1959; Biske and Devyatova, 1965; Heikki et al., 1995). The mollusc fauna listed in Table 1 is derived from borings on the south side of the watershed (Povenets) and from a number of isolated occur-
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PALAEO 2866 25-7-02 Cyaan Magenta Geel Zwart Fig. 5. Pollen diagram from boring in the city of Petrozavodsk, period of maximum marine inundation and inferred existence of the White Sea^Baltic seaway shaded. Pollen frequencies calculated according to Grichuk (1961).
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rences on the watershed up to altitudes of 96^101 m (Goretskyi, 1949; Biske, 1959; Devyatova and Loseva, 1964). With 24 species the faunas are signi¢cantly richer than those from nearby Petrozavodsk and St. Petersburg. Astarte sulcata, Trichotropis borealis, and Nucella lapillus show that warm saline Atlantic water from the White Sea £owed over the threshold and into the Onega basin. This is con¢rmed by the presence of Panomya arctica and N. lapillus at Povenets and saline diatoms at nearby Medvezhegorsk on the Onega side of the watershed. Also here the marine interval seems to be restricted to within the period from the Quercus^Pinus to the Taxus/Tilia zones (Znamenskaia and Cherminisova, 1962; Biske and Devyatova, 1965). During the maximum transgression the watershed located immediately to the north of Lake Onega at 105 m asl was penetrated by two 30^ 40-km-long and ca. 10-km-wide sounds (Fig. 1), which opened northwards into a wide shallow archipelago extending to the present White Sea shore (Atlas Karelskoi ASSR, 1989; Goretskyi, 1949; Apukhtin and Krasnov, 1966). This may be somewhat conjectural because the present watershed area is composed of Weichselian sediments. However, the faunal contrast between areas on and to the north of the watershed and those to its south show that during the Eemian a major hydrographical constriction of the seaway existed at this place. 2.8. Arkhangelsk region The Severnaya Dvina basin is the classical area for sediments from the Boreal Transgression. The basin contained a shallow marine inlet up to 500 km from the present river mouth at the White Sea. In the coastal area this basin was, via its tributary valleys, fused with the basins of the neighbouring Onega and Mezen rivers to form a large shallow archipelago extending from the Onega River basin to the Timan Ridge (Devyatova, 1982). Most of the area was ice-covered during the Weichselian (Larsen et al., 1999), but the Eemian sediments are mostly undisturbed and in situ. They are exposed in riverbanks and comprise
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every conceivable coastal and o¡shore habitat. Reviews of the very comprehensive literature dealing with the Eemian in this area were given by Lavrova (1937), Legkova (1967), and Devyatova (1982). In the coastal area to the north of Arkhangelsk pollen analysis from a boring at Izhma shows that the marine transgression reached the present coastline of the White Sea in the Late Saalian and marine conditions persisted into the Weichselian (Pleshivtseva, 1972). On the very southern margin of the basin, at Pasva, the marine inundation was restricted to the Quercus zone (Devyatova, 1982). Midway between these two localities, at Krasnaya Gorka, the transgression began in the Quercus zone and lasted into the Picea zone (Fig. 3 and Devyatova, 1982). The Eemian marine sediments begin with a lower shoreface facies. This is bounded below by a £ooding or ravinement surface, and followed by up to 30-m-thick o¡shore mud; followed again by shoreface and £uvial sediments. Further upriver shoreface sand dominates the sequences (Lavrova, 1937). The fauna listed in Table 1 is based on the lists by Zharkhidze (1963), Legkova and Schukin (1967), and our own observations, and comprises a wide range of habitats. This is to some extent the explanation for the unusual richness with 112 species, the richest among the regions presented here. The earliest transgression sediments contain a high number of oceanic species, such as Boreocingula castanea and Astarte sulcata as well as several Yoldiella species, showing that during this early phase even the most remote areas were oceanic or polyhaline. A transition to Macoma balthica-dominated faunas in the upper shoreface sediments at many localities may indicate that a change to estuarine conditions with reduced salinity took place at the start of the regression after the Quercus and Quercus^Corylus zones, as suggested by Devyatova (1982).
3. Geometry and bathymetry of the seaway The seaway was, according to our reconstruction, a ca. 150-km-wide sound extending from the
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North Sea to the south and west coast of the present White Sea, and via a broad connection further north to the Barents Sea, a distance of ca. 2500 km (Fig. 3). During the Holocene the three Danish straits, LillebZlt, StorebZlt and \resund, are the restrictive factor for water exchange between the Baltic and the ocean. In their present state these straits were shaped as £uvial valleys, eroded by rivers that drained the Baltic during its fresh water stages in the Late Weichselian and Early Holocene (Bjo«rck, 1995; Andre¤n et al., 2000). StorebZlt today carries 75% of the brackish outlet water from the Baltic and almost all in£ux of oceanic water from Kattegat (e.g. Segerstrafile, 1957), but there is no evidence to indicate that it existed in the Eemian. The same can be said for the narrow winding LillebZlt. Along \resund and in the interior of SjZlland, on the other hand, typical Eemian molluscs (Lucinella divaricata, Paphia senescens) occur scattered in Weichselian meltwater sediments and beach sediments (\dum, 1933; Madsen, 1965). Although sparse and circumstantial, this could indicate that a major entranceway to the Baltic traversed SjZlland. A candidate for such a waterway could be the buried Alnarp^Esrum Valley. This is a more than 100-km-long, more than 10-km-wide and more than 50-mdeep rami¢ed valley system crossing southern Sweden and northern SjZlland on its way from Kattegat to the Baltic (Miller, 1977; Konradi, 1992; Houmark-Nielsen, 1999). The valley system was shaped by faulting in the Palaeogene and Mesozoic substrate, and last functioned as a meltwater channel during the Weichselian when it was ¢lled with sand and gravel (Houmark-Nielsen, 1999). Although no Eemian sediments occur in the valley, the scattered ¢nds of reworked Eemian shells in its vicinity could indicate that it served as a major corridor for water exchange between the Baltic and Kattegat during the Eemian, thereby circumventing the thresholds at Darss and Drogden, which presently form a barrier for the entrance of dense oceanic water into the Baltic. As noted above a connection from the ocean to the Baltic may also have existed through Schleswig-Holstein in the area of the Kiel Canal, but the
outline of this waterway, a winding pattern of broads and narrow passages (Kosack and Lange, 1985), indicates that it was similar to the present Limfjord traversing northern Jutland without playing a signi¢cant role in the water exchange of the Baltic. In the north, from a much-enlarged White Sea the entrance gradually narrowed southwards over northern Karelia towards the major constriction at the watershed north of Lake Onega (Goretskyi, 1949; Atlas Karelskoi ASSR, 1989). The dimension of this entrance was in its dimensions similar to the present step-by-step entrance to the Baltic from the North Sea through Kattegat and the Danish straits, and both its outline and the faunal contrast between the areas to the north and south suggest that this was a major obstacle limiting the capacity for water transport through the seaway. A shallow side entrance from the Severnaya Dvina basin into Karelia may have existed along the river Voda (Devyatova, 1982). A peculiar feature of the Eemian coastline is its lobate appearance with the sea everywhere forming saline estuaries in present day valleys and depressions, contrasting to the present mature coastlines. This suggests rapid sea level rise and/or high tidal amplitude. Since this feature is found in all parts of the region rapid sea level rise may be the most likely explanation. The bathymetry of the seaway on Fig. 3 is based mainly on present depth contours to which are added the maximum altitude of the Eemian marine inundation, but no correction has been made for glacial erosion during the Weichselian when both the Baltic and the White Sea served major outlets for the Scandinavian ice sheet. As at present the Baltic basin was composed of a series of deep troughs with depths exceeding 200 m, separated by shallower thresholds. The same pattern was probably found in the Karelian passage where deep troughs occurred in the Ladoga and Onega depressions. The most signi¢cant deviation from the present bathymetry is the trough that may have extended across northern Jutland and into Kattegat, as indicated by borings showing great thickness of marine sediments of probable Eemian age (Lykke-Andersen et al., 1993; Seidenkrantz and Knudsen, 1997). The trough, which
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Table 3 Biogeographical and salinity grouping in Eemian faunas
apparently was ¢lled up during the Eemian, marks a signi¢cant deviation from the Holocene Kattegat, and may have played an important role in the hydrography. In summary, the White Sea^Baltic seaway was a ca. 150-km-wide sound extending for ca. 2500 km from the North Sea to the southern Barents Sea composed of more than 200-m-deep basins, separated by shallow thresholds. A major constriction existed in the north, to the north of Lake Onega, while a deep trench in Kattegat and a deep valley system extending through SjZlland and Skafine may have a¡orded a more e⁄cient water exchange than the Holocene. A channel through Schleswig-Holstein probably supplied restricted in£ow from the North Sea.
4. Duration The history of the seaway is a product of isostasy and eustasy following removal of the Saalian ice sheet, and at each site the duration of sea cover is determined by the interaction between these two forces. Since the palynological results come from the coastal fringes of the seaway, they can give only a minimum estimate of the duration of sea cover in each area.
The premises for pollen analytical dating of the marine sediments in each area have been discussed above, and the duration of marine inundation in each area is shown in Table 2 and Fig. 3. Outside the seaway, at the entranceways on the shore of the White Sea and in Kattegat borings have shown continuous sea cover from the Late Saalian through the Eemian and into the Weichselian (Pleshivtseva, 1972; Knudsen, 1994; Seidenkrantz and Knudsen, 1997; Kristensen et al., 1998). In the Wadden Sea marine invasion with warm fauna took place already in the Betula^Pinus zone, at the beginning of the Eemian. Recent results from Latvia indicate that the Baltic was also inundated with saline water at this time (Kalnina, 2001). Shortly afterwards, in the Pinus^Ulmus zone ca. 100 yr into the Eemian, Petrozavodsk was inundated, showing that the seaway into the Baltic was now opened to the Barents Sea. In£ux from the North Sea is documented ¢rst in the faunas of the Vistula valley from the beginning of the Quercus zone, 200 yr later. At this time also the northern parts of the Gulf of Bothnia were probably inundated. These results could indicate that the seaway opened ¢rst to the north and shortly afterwards to the west. However, as noted above, the pollen zones are not precisely
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Fig. 6. Inferred development of salinity in the southern Baltic during the Holocene and the Eemian. Salinity colours as in Fig. 3. The Holocene development is based on results from the Bornholm basin (Sohlenius et al., 2001).
synchronous in this large region, and are not suf¢ciently well de¢ned to distinguish events which are only a few hundred years apart, but it can be concluded that the opening of the Baltic to the Barents Sea and to the North Sea came in close succession within the ¢rst centuries of the Eemian. The regression began in the Corylus or Taxus/ Tilia zone (Petrozavodsk, Gulf of Bothnia). At many sites there is evidence for shallowing of the water depth in the Carpinus zone, and in the Picea zone the water retreated from many areas (Schleswig-Holstein, Vistula, Prangli). The records from Mga and the Ladoga shore show that that the Baltic was still in connection with the ocean at the end of the Eemian. From this, the Baltic was in continuous connection with the ocean at least from the beginning to the end of the Eemian, and there were, at least in the southern Baltic, no fresh water stages like the
Early Holocene Ancylus stage. The seaway from the North Sea to the White Sea functioned only in the Early Eemian from the Pinus^Ulmus or the Quercus zone to within the Taxus/Tilia zone, a period of 2^2.5 ka.
5. Hydrography and climate Table 3 summarises information on distribution, salinity and number of species recorded in each area. 5.1. Salinity and temperature In the Belt Sea the Cyprina clay contains a sparse fauna dominated by such small and short-lived species as Corbula gibba. In its structure the fauna is similar to the Abra alba bio-
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coenosis, which also today inhabits these areas (Thorson, 1957). This indicates that also in the Eemian this was an area of protected fjords with shallow wave base, large in£ux of terrestrial nutrients, and reduced salinity, as shown also by the frequent occurrence of Balanus improvisus. However, at the few localities where it has been preserved the shoreface Tapes or senescens sand may with its frequent Chamaelea gallina and Spisula elliptica compare to the C. gallina biocoenosis, which today inhabits sandy bottoms in the more saline Kattegat, but is absent from the Belt Sea and the Baltic (Thorson, 1957). The fauna in the scattered occurrences of Turritella clay is very similar to today’s Amphiura biocoenosis on deeper water in Kattegat (Thorson, 1957). This indicates that during the Eemian the salinity both in the upper and lower part of the water column in the Belt Sea and the western Baltic was as high as those of the present Kattegat, ca. 25x. This is supported by the absence or rarity of the Macoma balthica biocoenosis, which has inhabited the Baltic shallow water during the Holocene (Segerstrafile, 1957), and the presence of Ostrea and Venerupis senegalensis as well as other salinity demanding species in the Vistula valley indicating a surface salinity of ca. 25x, which is 15x higher than now and ca. 10x higher than the Holocene maximum. The evidence from Poland indicates that this salinity lasted from the Quercus zone and until regression set in at the end of the Carpinus zone (Makowska, 1986). In addition, the occurrence of Lusitanian species, notably Lucinella divaricata, in this area indicates winter sea surface temperatures at ca. 9‡C, 6‡C warmer than now. Fig. 6 compares the inferred salinity development during the Holocene and the Eemian in the southern Baltic. The Holocene is deduced from a record from the Bornholm basin, characterised by its short Ancylus stage and long and brackish initial Littorina stage (Sohlenius et al., 2001). The Early Eemian history was apparently simpler than that of the Holocene and lacks the repeated changes between fresh and brackish. We ascribe this to the dominance of eustatic sea level rise over isostatic rebound in the Eemian, as discussed above. Apart from this di¡erence the de-
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velopment seems to be in phase, with the salinity maximum being achieved in the Early Holocene and Eemian ^ only the Eemian Baltic was considerably more saline throughout. We explain this by a combination of factors as discussed below. In the White Sea area and Arkhangelsk region the situation was similar to the southern Baltic. The shoreface faunas, which now are subarctic, were dominated by such boreal species as Arctica islandica, Spisula elliptica and Cerastoderma edule, showing that salinities and temperatures were considerably higher than now, and the area was probably year-round ice-free. This lasted at least until the Carpinus zone, when regression began. In contrast, the low diverse faunas of Karelia and the St. Petersburg area, seven to eight species, re£ect as noted above low salinity and temperature and, in the St. Petersburg area, probably widespread anoxia. The dominating species, Portlandia arctica, is stenothermal and Jensen (1942) found that it requires bottom water temperatures below 4‡C, and preferably below 2.5‡C. Somewhat higher surface temperatures are indicated by the occurrence of Cerastoderma glaucum and Mytilus edulis. The Portlandia fauna has no analogue in the present Baltic, but similar biocoenoses exist in isolated bays with restricted water exchange along the coasts of the White Sea (Zenkevitch, 1963). In these bays the shallow waters are heated to high summer temperatures, as much as 20‡, and the shoreface fauna is composed of M. edulis, Macoma balthica and such boreal outliers as Littorina littorea and the plant Zostera marina, while the poorly oxygenated denser bottom water at 15 m depth with 6 0‡ temperatures houses an arctic P. arctica biocoenosis. During the Eemian the poor benthic faunas in the Karelian passage therefore point to long lasting water mass strati¢cation with low saline (ca. 15x) surface water and denser ( s 25x) cold (W2.5‡) bottom water with frequent anoxia. It should be noted that although P. arctica nowhere occurs as far south as St. Petersburg, the present faunas in the Baltic do have an arctic aspect, especially among crustaceans, as discussed intensively for more than a century (e.g. Segerstrafile, 1957), and Astarte borealis and Macoma calcarea, two species with their main distribution
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Fig. 7. Inferred surface hydrography of the Baltic at present (A) and Eemian (B). Colours for surface salinity as in Fig. 3. Arrows: red: warm surface current; blue: cold bottom current; green: brackish surface current (W10x); striped red/yellow: coastal water surface current ( s 15x); yellow: major source of river runo¡. Thickness of arrow signi¢es quantity. Present hydrography based on Segerstrafile, 1957; Zenkevitch, 1963; Kullenberg, 1981; Ehlin, 1981.
in the Arctic, thrive in the cold bottom water of the southern Baltic (Zenkevitch, 1963). The lack of other arctic species could possibly be a consequence of the Ancylus Lake stage, which in the Early Holocene interrupted the marine history of the Baltic (Fig. 6 and Bjo«rck, 1995; Andre¤n et al., 2000). Uninterrupted marine conditions going back to the Late Saalian may be a part of the explanation for the persistence of the Portlandia fauna at these southerly latitudes. 5.2. Hydrography and climate Fig. 7 compares the Early Eemian hydrography with that of the present. Conspicuous di¡erences are the larger size of the eastern Baltic, and the higher surface salinity of the southern Baltic in the Eemian. During the Holocene the restricted capacity of the Danish straits has been the main restriction for in£ow of saline water into the Baltic, and we suggest that the much higher salinity during the Eemian was partly a result of wider and deeper entranceways through Denmark. As
noted above the entrance between Norway and Denmark was wider, and a deep trench may have existed in Kattegat. Further, the Alnarp^Esrom valley could possibly have provided a short cut between Kattegat and the Baltic, and a more limited in£ow may have occurred through Schleswig-Holstein (Fig. 3). Other factors also played an important role. Thus the salinity in the recruiting area, the North Sea, was higher, as shown by the faunas in the Wadden Sea; and higher temperatures caused higher evaporation from the sea surface. Better mixing of the surface water, as indicated by the apparent absence of the Macoma balthica biocoenosis, may have been a¡orded by wind as at present and/or more vigorous tidal currents. Finally, the fresh water input was more dispersed. This applies especially to the Neva drainage, which today delivers ca. 20% of the Baltic fresh water input concentrated into the Gulf of Finland (Ehlin, 1981). In the Eemian this supply was distributed over and possibly caught in the Karelian basins. As noted above the faunas in Karelia and the
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St. Petersburg area suggest strati¢cation between low saline surface water and denser and colder bottom water with frequent anoxia. This implies very restricted advection of saline water into the Karelian passage and cold winters in this area, but during the period of the White Sea^Baltic seaway there was at least occasionally over£ow of saline surface water from the White Sea into the Lake Onega basin. This is shown by the saline fauna at Povenets. However, the contrast between this and the fauna from Petrozavodsk and the St. Petersburg area indicates that the saline in£ow both from the north and the south was insu⁄cient for ventilating the Karelian basin. In the St. Petersburg area the warm and saline £ora of planktonic diatoms may also imply £uxes of advective surface water from the southern Baltic, but it seems likely that a major restriction to the £ow must have existed also in the area of the Gulf of Finland. These results show that the White Sea^Baltic seaway during its 2^2.5-ka existence was not a major corridor for north^south transport of cold or warm water, and it is unlikely that its opening and closure had any signi¢cant in£uence on the north European climate. This is in contradiction to theoretical hypotheses expressed recently in a number of papers to explain changing oceanic and continental climates in northern Europe during the Eemian (Zagwijn, 1996; Kristensen et al., 1998; Bjo«rck et al., 2000; Rioual et al., 2001). Instead we suggest that these di¡erences may ¢nd their cause in the larger extent of the Eemian Baltic, and its more vigorous ventilation, both of which culminated in the Early Eemian, and gradually changed to regression and lower salinity.
6. Conclusions Our studies suggest that ^ the Baltic was connected with the ocean throughout the Eemian. In the Belt Sea and southern Baltic temperature and salinity were generally ca. 6‡C warmer and 15x higher than at present, culminating in the Early Eemian before the beginning of the Carpinus zone. After this the sea regressed and salinity declined.
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^ a seaway through Karelia connected the North Sea with the Barents Sea for 2^2.5 ka until isostatic uplift severed it at the watershed to the north of Lake Onega during the Taxus/Tilia zone or earlier. The Onega^Ladoga region continued to be an arm of the Baltic for several millennia, the Ladoga arm probably until the end of the Picea zone. ^ the seaway never functioned as a corridor for large-scale transport of water masses, and its opening and closure probably did not e¡ect the general ocean circulation or north European climate. ^ the higher salinity and more vigorous circulation in the Baltic and Belt Sea were caused by larger capacity of the Danish entranceways, higher salinity in the recruiting area of the North Sea, higher evaporation, and a more dispersed supply of river runo¡. ^ the passage between the present Baltic and White Sea, Karelia and the St. Petersburg region, had long lasting strati¢cation with brackish surface water and dense cold bottom water and frequent anoxia, similar to present conditions in the Gulf of Bothnia. ^ the contrast between the warm and well ventilated western and southern Baltic and the cold and stagnant water of Karelia and St. Petersburg implies that the Eemian oceanic to continental climate gradient from west to east over northern Europe was steeper than the Holocene.
Acknowledgements This study was carried out under the auspices of the ESF (European Science Foundation) administered QUEEN project (Quaternary environments, Eurasian Arctic), and the EU (European Union) Balteem project (Palaeoenvironmental and palaeoclimatic evolution of the Baltic Sea basin during the last (Eemian, Mikulino) interglacial). Fieldwork in Russia and Denmark was funded by the Danish Research Council’s grant to the CATLINA and CLIENT projects. All these projects provided fora for numerous and valuable discussions. We are especially indebted to Valery Astakhov, Dmitri Malakovskiy, and
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Alexei Matiouchkov for help and guidance. The artwork was performed by Lisa Bellhage, Louise Hansen, and Bent Knudsen. Very valuable comments and stimulating discussion were provided by Tom Meijer, and the two reviewers Matti Eronen and Richard Preece.
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