Early Holocene land floras and faunas from Edgeoya, eastern Svalbard

OLE BENNIKE and LARS HEDENAS

Bennike, 0. & Hedenas, L. 1995: Early Holocene land floras and faunas from Edge~ya,eastern Svalbard. Polar Research 14(2), 205-214. Early Holocene, near-shore marine sediments from Visdalen, Edgeaya, eastern Svalbard contain locally abundant allochthonous remains of land , notably bryophytes. Wetland species indicative of mineral- rich and calcareous soils are frequent, but upland plants are also well represented. The fossil assemblages are indicative of ecological and climatic conditions similar to those on Edge~yatoday. The sediments contain one of the first fossil beetles reported from Svalbard. Apparently, the modern flora of Svalbard was already established in the earliest Holocene, probably following immigration from northern Europe. A few Armeria scabra remains are believed to be derived from interglacial deposits. Ole Bennike, Geological Survey of Denmark and Greenland. Thoravej 8, DK-2400 Copenhagen NV, Denmark; Lars Hedenas, Department of Crvptogamic Botany, Swedish Museum of Natural History, Box 50007, $104 05 Stockholm, Sweden.

Europe demonstrate the mobility of plants (Birks Introduction 1994). Apart from exotic driftwood, pre-Holocene, Palaeoecological studies of Early Holocene Quaternary remains are rare in Svalbard terrestrial/limnic sediments have been carried out (Ingolfsson et al. 1995); however, a number of by Hyvarinen (1968, 1970, 1972). Surova et al. Holocene lake and peat deposits have been inves- (1982), Gottlich & Hornburg (1982), van der tigated, mostly by means of pollen analyses. Only Knaap (1989), Birks (1991) and Wohlfahrt et al. a few of these go back to the Early Holocene, (1995). In addition, there is a find of Polytrichum and very little is known about the immigration coinrnune Hedw. dated at 9870 * 140 years BP history of Svalbard flora and fauna. An exception (Salvigsen & Mangerud 1991). Surova et al. is the dyarf birch Betula nana that seems to have (1982) obtained a 14C date of 10,360 * 260 years immigrated to central Spitsbergen around 6000 BP (Tln-270) on basal peat from Reindalen on I4C years BP (Surova et al. 1982). Nathorst (1883) central Spitsbergen. They presented a rather was of the opinion that a few hardy species poss- crude pollen diagram from the peat section with ibly survived the last glacial stage on Svalbard. only few pollen and spore types from indigenous but that by far the majority of the vascular plants plants. The oldest peat date appears to come from immigrated after the last ice age via a land bridge Agardhdalen, western Spitsbergen, from where joining Svalbard and Scandinavia. Other scien- Punning et al. (1978) reported a date of tists, however, find support for the refugium 10.570 * 360 years BP (Tln-232). Van der Knaap hypothesis (R~nning1965; Odasz 1991; Odasz et (1989) obtained a I4C date of 6710 5 80 years BP al. 1991). One argument is that the plants could on basal peat from a section in Rosenbergdalen, not immigrate to Svalbard after the last ice age northwestern Edgeoya. Both macrofossil and pol- because no land bridge connected the archipelago len diagrams were published from the site. Only with the European mainland. However, the fact macrofossils of Salk polaris, Gramineae and five that the small. isolated, volcanic island Jan taxa were found in the section, but to this Mayen, which has never been connected to any must be added 25 pollen and spore types. Van continent by a land bridge, has been populated der Knaap concluded that the deposit and flora by 62 species of vascular plants (Lid 1964) shows of the former mire indicated that the climate was that long distance dispersal of plants does take warmer than today during peat accumulation. place across oceans in the . Many studies of Palaeoecological studies of lake deposits from full and late-glacial phytogeography in northwest Svalbard are even fewer than those of peat 206 Ole Bennike & Lars Hedenas deposits. Hyvarinen (1968. 1970. 1972) studied and animals that throw light on the history of lake sediments on northern Svalbard and Bj~rn- Svalbard flora and fauna. The remains come from Bya by means of pollen analysis. One of the sediment samples from Visdalen on the north- sequences from northern Svalbard dated back to western part of the island of Edgeoya, eastern ca 1Oka. but the interpretation of the pollen Svalbard (Fig. 1). A preliminary report on the diagrams is difficult due to a high content of present material that included some tentative long distance transported pollen. Birks (1991) moss identifications was published by Bennike presented a detailed macrofossil diagram from ( 1992). lake sediments from Skardtj~rna,western Spits- bergen. The basal sediments were dated by accel- erator mass spectrometry (AMS) on Salk and Study area Saxifraga remains to 8 ka. There is no safe indi- Edgeoya covers 5 150 km2 and is the largest island cation in Birks' study that any plants arrived after in the southeastern part of the Svalbard archi- this date. whereas three taxa were found that do pelago; extensive areas are glaciated, although not presently grow in the area. and a mean July the island is relatively low (Norderhaug 1970). temperature about 2°C higher than today was The bedrock is dominated by Triassic sandstone inferred for the Early Holocene. Wohlfahrt et al. and mudstone with layers of limestones and thin (1995) suggested that the mean July temperature coal seams in places, intruded by some dolerite on Bjsrnoya may have been as much as 45°C sills (Lock et al. 1978). The northwestern part of higher than present in the Early Holocene, based the island suppoits the richest vegetation and on beetle data. Terrestrial plant macrofossils in belongs to the Salix polaris zone of Brattbakk near-shore marine sediments from Adventfjor- (1986). Large areas are virtually barren, but the den, wchtern Spitsbergen. were studied by Heer dwarf Salixpolaris is common in favourable (1870) and Anderson (1910). They found abun- sites. Ninety-three species of vascular plants are dant stones of Empetrum tiigritm that no known from the island (Neilson 1970). From the Longer in this area. flora and vegetation a mean temperature for the The aim of this paper is to describe and inter- warmest month of around 2°C is suggested for the pret some remains of terrestrial and limnic plants northwestern part of the island.

big. 1. Left. Map of Sralbard. The arrow shows the location of Visdalen on Edgeoya. Right: Map of Visdalen ahowing thc location of the studied aampleb Contour interval 20 rn; contours only showed in Visdalen. For detailed description of the sites. see Moller er al. (19Y5) Early Holocene land floras and faunas from EdgeGya, eastern Soalbard 207

Turcz. We use the name A. scabra which is the Material and methods one currently in use in Greenlandic botany Seven samples were collected during the PONAM (Bocher et al. 1978). It is possible that some of 1991 expedition to eastern Svalbard. For the seeds referred to Draba sp. represent other descriptions and interpretations of the sampled Brassicaceae genera. All Salk remains (, sediments, see Moller et al. (1992, 1995). The bud scales, fruit capsules, bark and wood) are sediment samples consisted of fine sand and silt referred to Salix polaris, to which the diagnostic with allochthonous plant and animal remains, leaves belonged, and which is the only some of which were well preserved, indicating species presently growing on Edgeoya. Tar- short transport. The samples were wet-sieved axacum achenes were similar to Taraxacum bra- through 0.42 and 0.21 mm sieves, and the residue chyceras achenes which have smaller spines than left on the sieves was sorted under a dissecting T. arcticum (Trautv.) Dahlst. achenes, the only microscope. other Taraxacum species on Edge~yatoday; but Most of the bryophyte remains were well pre- if the achenes are interglacial in age the species served, which made it possible to identify a large identification becomes uncertain. proportion of the material to a taxonomic level Lepidurus was represented only by apodous where a partial reconstruction of the paleo- segments. For climatic reasons the remains are environment becomes possible. The bryophyte referred to Lepidurus arcticus. Olophrum was material is deposited in the Department of Cryp- represented by one elytron and possibly a tergite. togamic Botany, Swedish Museum of Natural His- The elytron can not be identified at the species tory (No. 520644-620650). The other material is level, but since 0. boreale is the only species housed in the Geological Museum, Copenhagen. recorded from Svalbard at present (Fjellberg The bryophyte nomenclature follows Corley et 1983), the fossil elytron has been referred to this al. (1981), with additions in Corley & Crundwell species (J. Bocher, pers. comm. 1992). (1991) except for Brachytheciurn coruscum I. Wag. (B. groenlandicq), Polytrichastrum alp- inum (Hedw.) G. L. Sm. (Polytrichum alpinum). P. sexangulare (Brid.) G. L. Sm. (Popytrichum Results and discussion sexangulare), Pseudocalliergon angustifoliurn Non - bryophytes Hedenas (Hedenas 1992b), P. trifarium (Web. & Mohr) Loeske (Calliergon trifarium), P. tur- Identified land and freshwater plants and animals gescens (T. Jens.) Loeske (Scorpidium turges- are listed in Tables 1 and 2 and some remains cens), Scorpidium cossoni (Schimp.) Hedenas are illustrated in Fig. 2. The samples were small (Drepanocladus cossoni) . Straminergon stram- (Table l), but nevertheless some of the assem- ineum (Brid.) Hedenas (Cal[iergon slramineum) blages are diverse. Samples 87711. 87728 and and Warnstofia sarmentosa (Wahlenb.) Hedenas) 86251 contained marine fossils, showing that these (Calliergon sarmentosum). Tracheophyte assemblages were deposited in shallow water mar- nomenclature follows Ronning (1979). ine environments, probably near the mouth of a river. The marine fossils comprise the following animal groups: foraminifera, hydroids, serpulids. ostracodes, barnacles, malacostracans, molluscs, Identification echinoids, bryozoans and fishes and the brown Seeds of Melandrium apetalum were identified algae Sphacelaria sp. and Laminaria sp. In this from their wing which is broader than that of the context it should also be noted that the oribatid related species M. angustiflorurn, but the seeds mite Ameronothrus lineatus, which occurred in only measured about 1.2 mm across. It is impos- similar samples not included in Table 1, lives at sible to make species identification of Armeria or near the sea shore (Hammer 1944). The other maritima s.1. calyces (Godwin 1975). but from assemblages derive from lagoon sediments phytogeographical considerations the taxon (88203) and glacio-lacustrine sediments (88651, represented must be one of the northern taxa 88655 and 86266) (Moller et al. 1995). that include Armeria arctica Wallr., A. maritima The radiocarbon dates range from 12.5 ka to (Mill.) Willd. subsp. arctica (Cham.) Hult., A. 8.5 ka (Table 3). The 12.5 ka date on sample scabra Pall.. A. labradorica Wallr. and A. sibirica 87711 does not date the time of sedimentation 208 Ole Bennike & Lars Hedenas

Tablr I Macrofossils (except brlophytes) from Edgeoya. Svalbard

Sample No. 87711 87728 88203 8625 I 86266 8865 I 88655 C-14 age (ka]' 12 5 >9 8 10 u 99 8.6 8.7 8.5 Localit! 1806 1715 1804 181 1 1810 2103 2103 Sample rire (liter) ns 0.35 0.1 07 1.0 0.2 1.1

PLANTS: Cenococcuni geoplirlum Fries + + ?Pyrenomycetidar + + Equrrerurn sp + + Ranun~iili~~sulphiirruJ Sol. - 1 Ranunculus sp - Papauer dalilrancitn Nordh + ++ Melandrrum aprrnluni Fenzl 1 3 Crrusrrunr arcricurn/aIpiniitn + ++ Minrinrrru rubella (Wbg.) Hiern + ++ Silene ucuulr (L.) Jacq + + Oxvrru dipria (L ) Hill ++ +it Pol~gonimiriviparuni L - + Arrnerra ,i.ahra Pall. - - Draba sp + + Salrx polarrs Wg + + Porenrillu \p + + Saxrfrugo opporrrrfolru L ++ ++ Dryas ocruperaln L - - Turnxaciim hrachycera Dahlrt - - Juncifi sp - + Lu;rrlu 5p + + Care.< \pp. + + ANIMALS. Lrpiduric\ urcricu$ Pallas Daphnra pder dc Gcer t!pe Candona \p. Oluphnrrn horrale (Pa! kull) 91chneumonidae Chironoinidae ?Lepidoptera Errgone 5p Orihalida -

+! A + i-+ Rrlatne frequent) 1-5 Absolute nurnberc 'The radiocarbon dates are presented in Table 3

since the sample is underlain by in situ shells may also be too young due to contamination by dated at 9.4ka (the sea water corrected age is intruding roots. 9.Oka) (Moller et al. 1995). The dated sample The assemblages are indicative of high arctic must therefore consist of older plus younger biotas similar to those on Edgeoya today. All material. Sample 87728 lies just under in situ vascular plants listed grow on Edgecbya today, shells dated to 9.7 ka (Table 3). The samples except one, namely Armeria scabru. which does with plant remains contained numerous small coal not occur on Svalbard at present. From its modern particles. and also a few pre-Quaternary trilete geographical range (Fig. 3), there seems to be no megaspores which could not be identified due to climatic reason for this since the plant grows in their simple morphology, but which are probably North Greenland and northernmost Canada of Mesozoic age (S. B. Manum, pers. comm. where the climate is similar to that of Svalbard. 1991). It is possible that previously published peat Because the radiocarbon date shows that the and gyttja dates based on bulk samples from the sample from the same location from which the region are too old due to contamination by coal Arnicria remains come includes some old plant particles. but on the other hand bulk peat samples material, it is possible that the Arrneria remains Early Holocene land floras and faunas from Edgelya, eastern Soalbard 209

Table 2. Bryophytes from Edgetiya. S = shoot; €3 = branch; L = .

Sample No. 87711 87728 88203 86251 86266 88651 88655

Amblystegium subg. Hygroamblystegium sp. Aulacomnium palustre A. turgidum Brachytheciurn coruscum B. turgidum Bryoeryrhrophyllum recuroirostrum Bryum cf. pseudotriquerrrtnr Bryum sp. Calliergon cf. richardsonii C. giganreurnlrichardson;; C. richarhonii Campylium polygamum Campylium sp Cerarodon purpureus Dichodontium pellucidum Distichium sp. Dirrichuni Pexicaule Drepanocladus s.str. sp. Encalypta alpinn E. proceralsrreptocarpa Encalypta sp. Isopterygiopsis pulchella Lophoria s.1. sp. Oncophorus wahlenbergii cf. Orthothecium sp, Paludella squarrosa Philonotis fontanalromentella Philonoris sp. Pogonarum dentaturn P. urnigerum Pohlia wahlenbergii Pohlia sp. Polytrichastrum alpinum P. sexangulare Polytrichum juniperinumlstricrum Polytricum sp. PolytrichumlPolyrrichastrum sp. Polytrichaceae sp Pseudocalliergon atigusrifolium P. turgescens Racomitrium canescenrlpanschii Racomitrium sp. Sanionia nioalis S. uncinara Scorpidium cossoni S. scorpioides Sphagnum squarrosumlreres Timmia austriaca T. noruegica Timmia sp. Tomenrypnunr nilens Warnstorfia sarmenrosa W. tundrae Indet. Indet. Amblystegiaceae Indet. acrocarp lndet. pleurocarp

*Of these, 18 and 5 shoots, respectwely, were found in coherent tufts '"Of these, 5 shoots were found in one coherent tuft. * * Talyptrae. 210 Ole Bennike & Lars Hedenas

Tuhle 3 AMS radiocarbon dates. For details of sediments and stratigraphv. see Moller el al. (1992. 1995).

Loc. Sample Altitude No No m asl. Lab No Age yrs BPI Dated material

1806 8771 1 19.8 AAR-846 12470 ? 250 Plant remains 1806 X77122 19.6 AAR-839 8960 ? 140 Mya truncata 1715 87721' 32.6 AAR-842 9710 ? 160 Mva truncatu 1x01 wo3 51.0 AAR-847 9965 ? 190 Plant remains 1811 86251 67.0 AAR-843 9475 ? 220 Plant remains4 2 103 8865 1 50.8 AAR-834 8690 -t_ 250 Plant remains 1610 86266 52.0 AAR-844 8640 ? 170 Plant remains 2103 88655 47.1 AAR-833 8530 -t 190 Plant remains

'The dates have been corrected for isotopic fractionation by normalising to b"C = -25% PDB. Samples 87712,87721, and 86251 have becn corrected for a marine reservoir age by subtracting 440 vears (Mangerud & Gulliksen 1975). 'Sample X7712 lies just below 87711 'Sample X7721 lies just above 87718. 'Remains of marine plants. are derived from interglacial sediments, in spite Wetland species are often common in fossil of the fact that they are well preserved. Armeria bryophyte assemblages (e.g., Janssens 1983; calyces are known from Last Interglacial deposits Dickson 1973. 1986; Hedenas 1992a; 1995) and in in central eastern Greenland (Bocher & Bennike the present samples Pseudocalliergon turgescens, 1991) and in northwestern Greenland (Bennike Scorpidium cossoni and Tomentypnum nitens, all & Bocher 1992): in addition, there is an undated occurring in mineral-rich to calcareous wetland northern extralimital occurrence from the eastern habitats today, were frequent. Other species of part of northern Greenland (Bennike unpubl.). more or less mineral-rich to calcareous wetlands The Turaxacum achenes are another possible include Brachythecium turgidum, Calliergon spp., interglacial candidate since none of the other Campylium polygarnurn, Campylium sp., Scor- samples contained this taxon: but as the plant pidium scorpioides. Warnsto@a tundrae and grows on Edge~yatoday. the achenes may also Pseudcalliergon angustifolium. The last species be Holocene. was recently described (Hedenas 1992b) and is The rove beetle Olophritm boreale has been known from only one extant Svalbard locality, on collected at Woodfjorden, northern Spitsbergen Bjorn~ya(Frisvoll & Hedenas unpubl.). In the and on Bjsrnoya (Fjellberg 1983) and may well Scandinavian mountain range it occurs mainly in be a member of the extant fauna of Edge~ya. moderate late snow beds in calcareous areas, but It has also been recorded from Holocene lake it occurs also in calcareous fens in the northern sediments on Bjsrn~ya(Wohlfahrt et al. 1995). boreal zone in Finland. This species is otherwise Olophrum boreale is a widely distributed northern known from Iceland, northern North America Holarctic species although it has not been found and the Chukotskiy Polustrov in easternmost Rus- in Greenland (Campbell 1983: Bocher 1988). sia (Hedenas unpubl.). Aulacomnium turgidum No limnic plants were found. but a few remains and A. palustre are also found in wetlands or of the limnic animals Lepidurus, Daphnia and on wet heaths, but these species are also found Candonu indicate that ponds with mesotrophic in intermediate (A. turgidurn) or mineral-poor waters existed in the area. habitats (A.palustre). Spring-influence or moving water is indicated by Paludella squarrosa, Phi- lonotis sp.. Pohlia wahlenbergii and Warnstorfia Brvophytes sarmentosa. All these taxa are found in inter- Many of the bryophytes (Table 2) indicate cal- mediate to rather mineral-poor places. Finally. cium- or mineral-rich conditions. This is what some of the taxa found indicate probable nutrient would be expected in a penglacial environment enrichment (N- and P- compounds), namely Cal- or in an area where the ice has recently retreated liergon s. str. spp. (i.e., excluding Pseudo- (cf. Miller 1987), but the bedrock of the area calliergon trifarium, Warnstorfia sarmentosa, probably also favours calciphiles. Straminergon stramineum), Campylium polyga- Early Holocene land floras and faunas from Edgelya, eastern Svalbard 21 1

Fig. 2. SEM photographs of selected macrofossils from Edgeoya. A-B. Armeria scabra calyces. C-D. Taraxacum brachycerar achenes. E. Melandrium apetalum seed. F. Ranunculus sulphurem achene. The figured specimens (MGUH 21272- 21677) are deposited in the type collection of the Geological Museum, Copenhagen. Scale bars = 1 mm.

mum, Drepanocladus s. str. sp., Sphagnum are either indifferent as regards the mineral squarrosumlteres and Warnstoijia tundrae (based content, or occur in relatively mineral-poor habi- on Kooijman 1993, van Wirdum 1991, Hedenas tats, whereas Dichodontium pellucidurn and Tim- & Kooijman unpubl. data, and own (Hedenas) mia norvegica indicate somewhat richer and extensive field-work in Northern Europe). wetter environments. Brachythecium coruscum, Many of the terrestrial taxa are found today in Bryoerythrophyllum recuruirostrum, Distichium more or less open (i.e., with sparse cover of sp., Ditrichum pexicaule, Encalypta spp., Isop- vascular plants) and unstable environments. terygiopsispulchella and Timmia austriaca all indi- Examples include Cerutodon purpureus, Pogon- cate mineral-rich habitats and can be found both atum dentatum, P. urnigerum, Polytrichastrum on somewhat disturbed soil (often in crevices or alpinum, P. sexangulare (often in late snow beds) when the soil surface is uneven) and in rock and Racomitrium canescenslpanschii. These taxa crevices in both dry and wetter situations. 212 Ole Bennike & Lars Hedeniis

large areas had a rather sparse cover of vascular plants. The upland plants may also (partly) reflect the environmental conditions some distance away from the wetlands. Possible means of transport include flowing water and wind. The assemblage is similar to what can be found in Svalbard (Kuc 1963; Philippi 1973; Elvebakk 1982) or at high altitudes on mineral-rich Scandinavian mountains today, and indicates that the climate was probably similar to that in these environments today.

Conclusions Many of the plant and animal taxa reported are new to the fossil flora and fauna of Svalbard, but this mostly reflects the few studies of Quaternary macroscopical terrestrial plant and animal ha 3. (‘ircumpolar map showing the modern geographical range of northern Armerra taxa (Armerraarcfica. A. scabra. A. remains that have been carried out. In addition, lubradoricu. A. sibiricu. A. maririma). based on HiltCn & Fries nearly all records are the oldest known from (1986) The arrow indicates the fossil locality. Svalbard. One of the results of this study is the possibility of finding interglacial plant remains on north- western Edgeaya, certainly not the most prom- ising area of Svalbard to search for such remains Saninnia uncinata occurs in a wide range of considering that pre-Holocene Quaternary habitats in northern and arctic areas. On the other deposits are almost unknown from this island hand, S. niualis is a species of extreme late snow- (Landvik et al. 1995). The data also show that the bed habitats or it may also grow along meltwater present flora, and the range of habitats and plant brooks from large late snow beds and glaciers. In communities of Svalbard, were already well estab- Europe. this species is at present known from lished in the earliest Holocene. We are inclined northern mountainous areas, where it is found to follow Nathorst’s view that only few vascular mainly in the middle and high alpine regions and plants survived the Weichselian glacial maximum in the Arctic, including Svalbard (Hedenas 1989). on Svalbard because most lowland areas were It is now aiso known from three northern North glaciated (Mangerud et al. 1992) and because the American and Greenland localities (Hedenas climate was colder than at present. If this is the unpubl.). but since the species was described case. the major part of the flora must have immi- rather recently (Hedenas 1989). andsince it seems grated in the latest part of the Weichselian. Many to be common in its northern European dis- radiocarbon dates between 13 and 10ka from tribution area. one would suspect that it is widely western Spitsbergen show that some areas were distributed in the Arctic. deglaciated by then (Mangrud et al. 1992). The find of Rmblysregiitrn subg. Hygroarn- Deglaciation of lowland areas on Edgeoya began hlwtegiitnz is interesting. The shoot resembles about 10.3 ka BP (Landvik et al. 1995). It should Atnblystegium fliwiatile. a species that is not very also be pointed out that virtually no clear endemic likely to be found in arctic areas. However. since species occur on Svalbard. a fact which also tes- the material does not permit identification to the tifies to the young age of the flora. species level. inferences regarding the habitat should not be made from this find. This shoot Acknowledgements. we thank C. Kronborg and 0.Stubdrup. rnav derive from interglacial deposits. University of Aarhus and P. Moller, Univcrsity of Lund. who Taken together. the bryophytes found indicate made the samples available for the analyses. J. Bocher, Zoologi- cal Museum, Denmark, kindly identified insect and spider mineral-rich environments, with a well-developed remains. and provided information on Ohphrum. J. Landvik wetland flora. The wetlands were probably sur- and two anonymous referees provided constructive criticism of rounded by somewhat unstable dry ground. where the manuscript Early Holocene land floras and faunas from Edge~ya,eastern Svalbard 213

Hyvarinen, H. 1968: Late-Quaternary sediment cores from References lakes on Bjorneya. Georgr. Ann. 50A. 235-245. Anderson. G. 1910: Die jetzige und fossile Quatarflora Spitz- Hyvarinen, H. 1970: Flandrian pollen diagrams from Svalbard. bergens als Zeugnis von Klimaanderungen. Pp. 409-417 in Geogr. Ann. 52A, 21S222. Posrglaziale Klimaueriinderungen. Stockholm. Hyvarinen, H. 1972: Pollen-analytical evidence for Flandrian Bennike, 0. 1992: Early Holocene terrestrial plants and animals climatic change in Svalbard. Pp. 225-237 in Vasari, Y., Hyva- from Edgeeya, eastern Svalbard. Lundqua Rep. 35,139-144. rinen, H. & Hicks, S. (eds.): Climatic changes in arctic Bennike, 0. & Bocher. J. 1992: Early Weichselian interstadial areas during the last ten-thousand years. Acra Unioersiraris land biotas at Thule, Northwest Greenland. Boreas 21, 111- Ouluenris, Series A, Scientiae Rerum Nacuralium No. 3, Geo- 117. Iogica No. 1. Birks. H. H. 1991: Holocene vegetational history and climatic HultBn, E. & Fries, M. 1986: Arlas of North European Vascular change in west Spitsbergen - plant macrofossils from Plants 1-111. 1172 pp. Koeltz Scientific Books, Konigstein. Skardtjarna. an Arctic lake. The Holocene I, 2W218. Ing6lfsson. 0..Rognvaldsson, F.,Bergsten, H., Hedenas. L.. Birks, H. H. 1994: Plant macrofossils and the nunatak theory Lemdahl. G. & Lirio, J. M. 1995: Late Quaternary glacial- of per-glacial survival. Pp. 12F143 in Lotter, A. F. & and environmental history of Kongsoya, Svalbard. Polar Res. Ammann. B. (eds.): Festschrift Gerhard Lang. Dissertariones 14(2), 123-139. Boranicce 234. Janssens, J. A. 1983: Past and extant distribution of Dre- Bocher, J. 1988: The Coleoptera of Greenland. Medd. Grpn- panocladus in North America, with notes on the dif- land Biosci. 26. 100 pp. ferentiation of fossil fragments. J. Hattori Bot. Lab. 54.251- Bocher, J. & Bennike, 0. 1991: Interglacial land biotas of 292. Jameson Land, East Greenland. Lundqua Rep. 33, 129- Kooijman, A. M. 1993: On the ecologlcal amplitude of four 136. mire bryophytes; a reciprocal transplant experiment. Lmf- Bocher. T. W., Fredskild, B., Holmen, K. & Jacobsen, K. bergia 18, 1S24. 1978: [email protected]. Haase & sen, Copenhagen. 327 pp. Kuc. M. 1963: Flora of and their distribution on the Brattbakk, I. 1986: Vegetatjonsregioner - Sualbard og Jan north coast of Hornsund (S.W. - Svalbard). Fragm. flor. Maven. Statens Kartverk, Hdnefoss (map). geobot. Y. 291-366. Campbell, J. M. 1983: A revision of the North American Landvik, J. Y.,Hjort, C., Mangerud, J.. Moller, P. Salvigsen, Omaliinae (Coleoptera: Staphylinidae). The genus 0. 1995: The Quaternary record of eastern Svalbard - an Olophrum Erichson. Can. Ent. 115. 577-622. overview. Polar Res. 14(2). 95-103. Corley, M. F. V. & Crundwell, A. C. 1991: Additions and Lid, J. 1964: The flora of Jan Mayen. Norsk Polarins!. Skr. 130. amendments to the mosses of Europe and the Azores. J. 107 pp. Bryol. 16, 337-356. Lock, B. E., Pickton, C. A. G., Smith, D. G.. Batten. D. corley. M. F. V., Crundwell. A. C., Dull, R., Hill, M. 0. & J. & Harland. W. B. 1978: The Geology of Edgeclya and Smith, A. J. E. 1981: Mosses of Europe and the Azores; an Barentseya, Svalbard. Norsk Polarinsr. Skr. 168. 64 pp. annotated list of species, with synonyms from the recent Mangerud, I. & Gulliksen, S. 1975: Apparent radiocarbon litterature. J. Bryol. 11, 609-689. ages of marine shells from . Spitsbergen. and Arctic Dickson. J. H. 1973: Bryophyres of the Pleistocene. Cambridge Canada. Quat. Res. 5, 26?-273. University Press. Cambridge. 256 pp. Mangerud, J., Bolstad, M., Elgersma, A., Helliksen, D.. Land- Dickson, J. H. 1986: Bryophyte analysis. Pp. 627-643 in vik, J. Y..Lenne, I.. Lycke, A. K., Salvigsen, 0..Sandahl. Berglund, B. E. (ed.): Handbook of Holocene Palaeoecology T. & Svendsen. I. I. 1992: The last glacial maximum on and Palaeohydrology . John Wiley & Sons, Chichester. Spitsbergen, Svalbard. Quar. Res. 38. 1-31. Elvebakk. A. 1982: Geological preferences among Svalbard Miller. N. G. 1987: Late Quaternary fossil mossflorasof eastern plants. Inter-Nord 16, 11-31. North America: evidence of major floristic changes during Fjellberg, A. 1983: Three species of staphylinid Coleoptera new the late Pleistocene-early Holocene transition. Svmp. Biol. to Spitsbergen. Fauna Noruegica Serie B 30, 110-111. Hungarica 35, 34S360. Godwin, H. 1975: The hisrory of the Bririrh porn. Cambridge Moller. P., Kronborg, C.. JohdnSSOn. K. & Stubdrup. 0. P. University Press, Cambridge. 541 pp. 2nd ed. 1992: The Quaternary history of Visdalen, Edgeaya. eastern Gottlich, K. & Hornburg, P. 1982: Ein Zeuge warmezeitlicher Svalbard. Lundqua Rep. 35, 85-130. Moore im Adventdalen auf Spitsbergen (Svalbard-Archipel). Moller, P., Stubdrup, 0. P. & Kronborg, C. 1995: Late Weich- TELMA 12, 253-260. selian to early Holocene sedimentatin in a steep fjord/valley Hammer, M. 1944: Studies on the Oribatids and Collemboles setting. Visdalen, Edge~ya, eastern Svalbard: glacial of Greenland. Medd. Grpnland 141 (3). 210 pp. deposits, alluvial/colluvial-fan deltas and spit-platforms. Hedenas, L. 1989: The genus Sanionia (Musci) in northwestern Polar Res. 14(2), 181-203. Europe, a taxonomic revision. Ann. Bot. Fennici 26, 399- Nathorst, A. G. 1883: Nya bidrag till kannedomen om Spets- 419. bergens karlvkxter, och deres vaxtgeografiska forhillan- Hedenas, L. 1992a ["1990']: A subfossil find of Pseu- den. Kgl. Suenska Verensk. - Akad. Handl., Ny foljd 20(61. docalliergon breuifolium in northern Sweden. Lindbergia 16, 88 PP. 150-152. Neilson, A. H. 1970: Vascular plants of Edgeeya, Svalbard. Hedenas, L. 1992b ["1990"]: The genus Pseudocalliergon in Norsk Polarinst. Skr. 150. 71 pp. northern Europe. Lindbergia 16, 80-99. Norderhaug. M. 1970: Investigations of the Svalbard reindeer Hedenas, L. 1995 y1994"l: Environments indicated by bry- (Rangifer rarandur platyrhynchur) in Barentsdya and ophytes in early Weichselian interstadial deposits from north- Edgeeya, summer 1969. Norsk Polarinsf. Arbok 1969. 70- ern Sweden. Lindbergia 19, 87-105. 79. Heer, 0. 1870: Die miocane Flora und Fauna Spitzbergens. K. Odasz, A. M. 1991: Distribution and ecology of herbaceous Suenska Vet. Akad. Handl. 8 (7). 23-24. Pedicularis dasynrha (Scrophulariaceae) in Spitsbergen, Sval- 214 Ole Bennike & Lars Hedenas

bard archipelago: relation to present and past environments Surova, T. G.. Troickij. L. S. & Punning, Ja.-M. K. 1982: Striae 34. 145-152 Ob istorii oledenenija Spicbergena v golocene PO dannym Odasz. A. M.,Karkkainen. K., Muona. 0. & Wein. G. 1991: paleobotaniEeskich issledovanij (On the history of the Hol- Genetic distances between geographically isolated Pediculark ocene glaciation of Spitsbergen in the Holocene from data of dasyantha populations in Spitsbergen. Svalbard Archipelago. palaeobotanical investigations). Marerialy gljaciologifeskich Noway. Evidence of glacial survival? Norsk Geol. Tidssk hsledouonjj. Chronika, obsuidsniju (Materials of glaci- 71. 21%221. ological investigations. Chronicle, Discussions) 42, 100-106 Philippi. G. 1973: Moosflora und Moosvegetation des Freeman- Van der Knaap. W. 0. 1989: Past vegetation and reindeer on Sund-Gebietes (Shdost-Spitsbergen). Ergebnirsc der Stuuf- Edgeeya (Spitsbergen) between c. 7900 and c. 3800 BP, stud- erland-Expedition 7. 83 pp. ied by means of peat layers and reindeer faecal pellets. J. Punning, J. M.. Rajamae, R. & Ehrenpreis. M. 1978: Tallinn Biogeogr. 16, 379-395. radiocarbon dates V. Radiocarbon 20. 4614. Van Wirdum. G. 1991: Vegetation and hvdrology of flootirig R~nning.0. I. 1965: Et bidrag ti1 forstielsen av de arktiske rich-fern. Datawyse, Maastricht. planters historie pi Svalbard Trondheim, Aktietrykkeriet 1 Wohlfahrt, B., Lemdahl, G.,Olsson, S., Persson, T., Snowball, Trondheim. 20 pp. I. & Jones, V. 1995: Early Holocene environment on Bjar- Rsnning. 0 1. 1979. Soulbards Flora. Norsk Polarinstitutr. n~ya(Svalbard) inferred from multidisciplinary lake sediment Oslo. 128 pp studies. Polar Res. 14(2). 253-275. Salvigsen. 0. & Mangerud. J. 1991: Holocene shoreline dis- placement at Agardhbukta. eastern Spitsbergen. Svalbard Polar Res. 9. 1-7.