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 E d g e ~ y a , 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 plants, 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 E d g e ~ y a today. 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. Introduction Apart from exotic driftwood, pre-Holocene, Quaternary plant remains are rare in Svalbard (Ingolfsson et al. 1995); however, a number of Holocene lake and peat deposits have been inves- tigated, mostly by means of pollen analyses. Only a few of these go back to the Early Holocene, and very little is known about the immigration history of Svalbard flora and fauna. An exception is the dyarf birch Betula nana that seems to have immigrated to central Spitsbergen around 6000 I4C years BP (Surova et al. 1982). Nathorst (1883) was of the opinion that a few hardy species poss- ibly survived the last glacial stage on Svalbard. but that by far the majority of the vascular plants immigrated after the last ice age via a land bridge joining Svalbard and Scandinavia. Other scien- tists, however, find support for the refugium hypothesis ( R ~ n n i n g 1965; Odasz 1991; Odasz et al. 1991). One argument is that the plants could not immigrate to Svalbard after the last ice age because no land bridge connected the archipelago with the European mainland. However, the fact that the small. isolated, volcanic island Jan Mayen, which has never been connected to any continent by a land bridge, has been populated by 62 species of vascular plants (Lid 1964) shows that long distance dispersal of plants does take place across oceans in the Arctic. Many studies of full and late-glacial phytogeography in northwest Europe demonstrate the mobility of plants (Birks 1994). Palaeoecological studies of Early Holocene terrestrial/limnic sediments have been carried out by Hyvarinen (1968, 1970, 1972). Surova et al. (1982), Gottlich & Hornburg (1982), van der Knaap (1989), Birks (1991) and Wohlfahrt et al. (1995). In addition, there is a find of Polytrichum coinrnune Hedw. dated at 9870 * 140 years BP (Salvigsen & Mangerud 1991). Surova et al. (1982) obtained a 14C date of 10,360 * 260 years BP (Tln-270) on basal peat from Reindalen on central Spitsbergen. They presented a rather crude pollen diagram from the peat section with only few pollen and spore types from indigenous plants. The oldest peat date appears to come from Agardhdalen, western Spitsbergen, from where Punning et al. (1978) reported a date of 10.570 * 360 years BP (Tln-232). Van der Knaap (1989) obtained a I4C date of 6710 5 80 years B P on basal peat from a section in Rosenbergdalen, northwestern Edgeoya. Both macrofossil and pol- len diagrams were published from the site. Only macrofossils of S a l k polaris, Gramineae and five moss taxa were found in the section, but to this must be added 25 pollen and spore types. Van der Knaap concluded that the deposit and flora of the former mire indicated that the climate was warmer than today during peat accumulation. Palaeoecological studies of lake deposits from Svalbard are even fewer than those of peat 206 O l e Bennike & L a r s Hedenas deposits. Hyvarinen (1968. 1970. 1972) studied lake sediments on northern Svalbard and B j ~ r n - Bya by means of pollen analysis. O n e of the sequences from northern Svalbard dated back to ca 1Oka. but the interpretation of the pollen diagrams is difficult d u e to a high content of long distance transported pollen. Birks (1991) presented a detailed macrofossil diagram from lake sediments from S k a r d t j ~ r n a , western Spits- bergen. The basal sediments were dated by accel- erator mass spectrometry ( A M S ) o n S a l k and Saxifraga remains to 8 ka. T h e r e is no safe indi- cation i n Birks' study that any plants arrived after this date. whereas three taxa were found that d o not presently grow in the area. and a mean July temperature about 2°C higher than today was inferred for the Early Holocene. Wohlfahrt et al. (1995) suggested that the mean July temperature on Bjsrnoya may have been as much as 4 5 ° C higher than present in the Early Holocene, based on beetle data. Terrestrial plant macrofossils in near-shore marine sediments from Adventfjor- den, wchtern Spitsbergen. were studied by Heer (1870) and A n d e r s o n (1910). They found abun- dant fruit stones o f E m p e t r u m tiigritm that no Longer fruits in this a r e a . The aim of this paper is to describe and inter- pret some remains of terrestrial and limnic plants and animals that throw light on the history of Svalbard flora and fauna. T h e remains come from sediment samples from Visdalen on the north- western part of the island of Edgeoya, eastern Svalbard (Fig. 1). A preliminary report on t h e present material that included some tentative moss identifications was published by Bennike ( 1992). Study area Edgeoya covers 5 150 km2 a n d is the largest island in the southeastern part of the Svalbard archi- pelago; extensive areas are glaciated, although the island is relatively low (Norderhaug 1970). T h e bedrock is dominated by Triassic sandstone and mudstone with layers of limestones and thin coal seams in places, intruded by some dolerite sills (Lock e t al. 1978). T h e northwestern part of the island suppoits the richest vegetation and belongs to the Salix polaris zone of Brattbakk (1986). Large areas a r e virtually barren, but the dwarf shrub S a l i x p o l a r i s is common in favourable sites. Ninety-three species of vascular plants a r e known from t h e island (Neilson 1970). From the flora and vegetation a mean temperature for the warmest month of around 2°C is suggested for the northwestern part of the island. big. 1 . Left. Map of Sralbard. The a r r o w 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. ( 1 9 Y 5 ) Early Holocene land floras and faunas from EdgeGya, eastern Soalbard 207 Material and methods Seven samples were collected during the PONAM 1991 expedition to eastern Svalbard. For descriptions and interpretations of the sampled sediments, see Moller et al. (1992, 1995). The sediment samples consisted of fine sand and silt with allochthonous plant and animal remains, some of which were well preserved, indicating short transport. The samples were wet-sieved through 0.42 and 0.21 mm sieves, and the residue left on the sieves was sorted under a dissecting microscope. Most of the bryophyte remains were well pre- served, which made it possible to identify a large proportion of the material to a taxonomic level where a partial reconstruction of the paleo- environment becomes possible. The bryophyte material is deposited in the Department of Cryp- togamic Botany, Swedish Museum of Natural His- tory (No. 520644-620650). The other material is housed in the Geological Museum, Copenhagen. The bryophyte nomenclature follows Corley et al. (1981), with additions in Corley & Crundwell (1991) except for Brachytheciurn coruscum I . Wag. ( B . g r o e n l a n d i c q ) , Polytrichastrum alp- inum (Hedw.) G . L . Sm. (Polytrichum alpinum). P . sexangulare (Brid.) G. L. Sm. (Popytrichum sexangulare), Pseudocalliergon angustifoliurn Hedenas (Hedenas 1992b), P . trifarium (Web. & Mohr) Loeske (Calliergon trifarium), P . tur- gescens (T. Jens.) Loeske (Scorpidium turges- c e n s ) , Scorpidium cossoni (Schimp.) Hedenas (Drepanocladus cossoni) . Straminergon stram- ineum (Brid.) Hedenas (Cal[iergon slramineum) and Warnstofia sarmentosa (Wahlenb.) Hedenas) (Calliergon sarmentosum). Tracheophyte nomenclature follows Ronning (1979). Identification Seeds of Melandrium apetalum were identified from their wing which is broader than that of the related species M . angustiflorurn, but the seeds only measured about 1.2 mm across. It is impos- sible to make species identification of Armeria maritima s.1. calyces (Godwin 1975). but from phytogeographical considerations the taxon represented must be one of the northern taxa that include Armeria arctica Wallr., A . maritima (Mill.) Willd. subsp. arctica (Cham.) Hult., A . scabra Pall.. A . labradorica Wallr. and A . sibirica Turcz. We use the name A . scabra which is the one currently in use in Greenlandic botany (Bocher et al. 1978). It is possible that some of the seeds referred to Draba sp. represent other Brassicaceae genera. All S a l k remains (leaves, bud scales, fruit capsules, bark and wood) are referred t o Salix polaris, t o which the diagnostic leaves belonged, and which is the only willow species presently growing on Edgeoya. Tar- axacum achenes were similar to Taraxacum bra- chyceras achenes which have smaller spines than T . arcticum (Trautv.) Dahlst. achenes, the only other Taraxacum species on E d g e ~ y a today; but if the achenes are interglacial in age the species identification becomes uncertain. Lepidurus was represented only by apodous segments. For climatic reasons the remains are referred to Lepidurus arcticus. Olophrum was represented by one elytron and possibly a tergite. The elytron can not be identified at the species level, but since 0. boreale is the only species recorded from Svalbard at present (Fjellberg 1983), the fossil elytron has been referred to this species (J. Bocher, pers. comm. 1992). Results and discussion Non - bryophytes Identified land and freshwater plants and animals are listed in Tables 1 and 2 and some remains are illustrated in Fig. 2. The samples were small (Table l ) , but nevertheless some of the assem- blages are diverse. Samples 87711. 87728 and 86251 contained marine fossils, showing that these assemblages were deposited in shallow water mar- ine environments, probably near the mouth of a river. The marine fossils comprise the following animal groups: foraminifera, hydroids, serpulids. ostracodes, barnacles, malacostracans, molluscs, echinoids, bryozoans and fishes and the brown algae Sphacelaria sp. and Laminaria sp. I n this context it should also be noted that the oribatid mite Ameronothrus lineatus, which occurred in similar samples not included in Table 1, lives at or near the sea shore (Hammer 1944). The other assemblages derive from lagoon sediments (88203) and glacio-lacustrine sediments (88651, 88655 and 86266) (Moller et al. 1995). The radiocarbon dates range from 12.5 ka to 8.5 ka (Table 3). The 12.5 ka date on sample 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 ( k a ] ' Localit! Sample rire (liter) PLANTS: Cenococcuni geoplirlum Fries ?Pyrenomycetidar Equrrerurn sp R a n u n ~ i i l i ~ ~ sulphiirruJ Sol. Ranunculus sp Papauer dalilrancitn Nordh Melandrrum aprrnluni Fenzl Crrusrrunr arcricurn/aIpiniitn Minrinrrru rubella ( W b g . ) Hiern Silene ucuulr ( L . ) Jacq O x v r r u d i p r i a ( L ) Hill P o l ~ g o n i m iriviparuni L Arrnerra ,i.ahra Pall. Draba sp Salrx polarrs Wg Porenrillu \p Saxrfrugo opporrrrfolru L Dryas ocruperaln L Turnxaciim h r a c h y c e r a Dahlrt J u n c i f i sp Care.< \pp. ANIMALS. Lrpiduric\ urcricu$ Pallas Daphnra p d e r dc Gcer t!pe Oluphnrrn horrale (Pa! kull) 91chneumonidae Chironoinidae ?Lepidoptera Errgone 5p Orihalida Lu;rrlu 5p Candona \p. 12 5 >9 8 10 u 9 9 1806 1715 1804 181 1 n s 0.35 0.1 0 7 8.6 8.7 8.5 1810 2103 2103 1.0 0.2 1.1 + + + + + + - 1 - + + + 1 3 + + + + + + + + + + + i t - + - - + + + + + + + + + + - - - - - + + + + + - +! A + i - + R r l a t n e frequent) 1-5 Absolute nurnberc 'The radiocarbon dates are presented in Table 3 since the sample is underlain by in situ shells dated at 9 . 4 k a (the sea water corrected age is 9.Oka) (Moller et al. 1995). T h e dated sample must therefore consist of older plus younger material. Sample 87728 lies just under in situ shells dated t o 9.7 ka (Table 3 ) . T h e samples with plant remains contained numerous small coal particles. and also a few pre-Quaternary trilete megaspores which could not b e identified due to their simple morphology, but which are probably of Mesozoic age ( S . B . M a n u m , pers. comm. 1991). It is possible that previously published peat and gyttja dates based o n bulk samples from t h e region are t o o old d u e to contamination by coal particles. but o n the other hand bulk peat samples may also be t o o young d u e to contamination by intruding roots. T h e assemblages are indicative of high arctic biotas similar t o those on Edgeoya today. All vascular plants listed grow on Edgecbya today, except o n e , namely Armeria scabru. which does not occur o n Svalbard a t present. From its modern geographical range (Fig. 3), there seems to be n o climatic reason for this since t h e plant grows in North Greenland and northernmost Canada where the climate is similar to that of Svalbard. Because t h e radiocarbon date shows that the sample from the same location from which the Arnicria remains come includes some old plant material, it is possible that the Arrneria remains Early Holocene land floras and faunas f r o m E d g e l y a , eastern Soalbard 209 Table 2. Bryophytes from Edgetiya. S = shoot; €3 = branch; L = leaf. Sample No. 87711 87728 88203 86251 86266 88651 88655 Amblystegium subg. 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. s p . Encalypta alpinn E. proceralsrreptocarpa Encalypta sp. Isopterygiopsis pulchella Lophoria s.1. sp. Oncophorus wahlenbergii cf. Orthothecium sp, Paludella squarrosa Philonotis fontanalromentella Philonoris s p . Pogonarum dentaturn P. urnigerum Pohlia wahlenbergii Pohlia sp. Polytrichastrum alpinum P. sexangulare Polytrichum juniperinumlstricrum Polytricum sp. PolytrichumlPolyrrichastrum sp. Polytrichaceae s p 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 Hygroamblystegium s p . *Of these, 18 and 5 shoots, respectwely, were found in coherent tufts '"Of these, 5 shoots were found in one coherent tuft. * * T a l y p t r a e . 210 Ole Bennike & Lars Hedenas Tuhle 3 A M S radiocarbon dates. For details of sediments and stratigraphv. see Moller e l al. (1992. 1995). Loc. Sample Altitude N o No m a s l . Lab No Age yrs BPI Dated material 12470 ? 250 Plant remains 1806 8771 1 19.8 AAR-846 1806 X77122 19.6 AAR-839 8960 ? 140 Mya truncata 1715 87721' 32.6 AAR-842 9710 ? 160 Mva truncatu 1x01 w o 3 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% P D B . Samples 87712,87721, and 86251 have becn corrected f o r 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 of the fact that they are well preserved. Armeria calyces are known from Last Interglacial deposits i n central eastern Greenland (Bocher & Bennike 1991) and in northwestern Greenland (Bennike & Bocher 1992): in addition, there is an undated northern extralimital occurrence from the eastern part of northern Greenland (Bennike unpubl.). The Turaxacum achenes are another possible interglacial candidate since none of the other samples contained this taxon: but as the plant grows on E d g e ~ y a today. the achenes may also be Holocene. The rove beetle Olophritm boreale has been collected at Woodfjorden, northern Spitsbergen and on Bjsrnoya (Fjellberg 1983) and may well be a member of the extant fauna of E d g e ~ y a . It has also been recorded from Holocene lake sediments on B j s r n ~ y a (Wohlfahrt et al. 1995). Olophrum boreale is a widely distributed northern Holarctic species although it has not been found in Greenland (Campbell 1983: Bocher 1988). No limnic plants were found. but a few remains of the limnic animals Lepidurus, Daphnia and Candonu indicate that ponds w i t h mesotrophic waters existed in the area. Brvophytes Many of the bryophytes (Table 2 ) indicate cal- cium- or mineral-rich conditions. This is what would be expected in a penglacial environment or in an area where the ice has recently retreated (cf. Miller 1987), but the bedrock of the area probably also favours calciphiles. Wetland species are often common in fossil bryophyte assemblages (e.g., Janssens 1983; Dickson 1973. 1986; Hedenas 1992a; 1995) and in the present samples Pseudocalliergon turgescens, Scorpidium cossoni and Tomentypnum nitens, all occurring in mineral-rich t o calcareous wetland habitats today, were frequent. Other species of more or less mineral-rich to calcareous wetlands include Brachythecium turgidum, Calliergon spp., Campylium polygarnurn, Campylium s p . , Scor- pidium scorpioides. Warnsto@a tundrae and Pseudcalliergon angustifolium. The last species was recently described (Hedenas 1992b) and is known from only one extant Svalbard locality, on B j o r n ~ y a (Frisvoll & Hedenas unpubl.). In the Scandinavian mountain range i t occurs mainly in moderate late snow beds in calcareous areas, but it occurs also in calcareous fens in the northern boreal zone in Finland. This species is otherwise known from Iceland, northern North America and the Chukotskiy Polustrov in easternmost Rus- sia (Hedenas unpubl.). Aulacomnium turgidum and A . palustre are also found in wetlands or on wet heaths, but these species are also found in intermediate ( A . turgidurn) or mineral-poor habitats ( A . palustre). Spring-influence or moving water is indicated by Paludella squarrosa, Phi- lonotis sp.. Pohlia wahlenbergii and Warnstorfia sarmentosa. All these taxa are found in inter- mediate to rather mineral-poor places. Finally. some of the taxa found indicate probable nutrient enrichment (N- and P- compounds), namely Cal- liergon s. str. spp. (i.e., excluding Pseudo- calliergon trifarium, Warnstorfia sarmentosa, Straminergon stramineum), Campylium polyga- Early Holocene land floras and faunas f r o m Edgelya, eastern Svalbard 21 1 Fig. 2 . SEM photographs of selected vascular plant 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 m m . m u m , Drepanocladus s . str. sp., Sphagnum squarrosumlteres and Warnstoijia tundrae (based on Kooijman 1993, van Wirdum 1991, Hedenas & Kooijman unpubl. data, and own (Hedenas) extensive field-work in Northern Europe). Many of the terrestrial taxa are found today in more or less open (i.e., with sparse cover of vascular plants) and unstable environments. Examples include Cerutodon purpureus, Pogon- atum dentatum, P . urnigerum, Polytrichastrum alpinum, P . sexangulare (often in late snow beds) and Racomitrium canescenslpanschii. These taxa are either indifferent as regards the mineral content, or occur in relatively mineral-poor habi- tats, whereas Dichodontium pellucidurn and Tim- mia norvegica indicate somewhat richer and wetter environments. Brachythecium coruscum, Bryoerythrophyllum recuruirostrum, Distichium sp., Ditrichum pexicaule, Encalypta spp., Isop- terygiopsispulchella and Timmia austriaca all indi- cate mineral-rich habitats and can be found both on somewhat disturbed soil (often in crevices or when the soil surface is uneven) and in rock crevices in both dry and wetter situations. 212 Ole Bennike & Lars Hedeniis ha 3. (‘ircumpolar map showing the modern geographical range of northern Armerra taxa (Armerra arcfica. A . scabra. A . lubradoricu. A . sibiricu. A . maririma). based on HiltCn & Fries (1986) The arrow indicates the fossil locality. Saninnia uncinata occurs in a wide range of habitats in northern and arctic areas. On the other hand, S. niualis is a species of extreme late snow- bed habitats or it may also grow along meltwater brooks from large late snow beds and glaciers. In Europe. this species is at present known from northern mountainous areas, where i t is found mainly in the middle and high alpine regions and in the Arctic, including Svalbard (Hedenas 1989). I t is now aiso known from three northern North American and Greenland localities (Hedenas unpubl.). but since the species was described rather recently (Hedenas 1989). andsince it seems to be common i n its northern European dis- tribution area. one would suspect that i t is widely distributed in the Arctic. The find of Rmblysregiitrn subg. Hygroarn- hlwtegiitnz is interesting. The shoot resembles Atnblystegium fliwiatile. a species that is not very likely to be found in arctic areas. However. since the material does not permit identification to the species level. inferences regarding the habitat should not be made from this find. This shoot rnav derive from interglacial deposits. Taken together. the bryophytes found indicate mineral-rich environments, with a well-developed wetland flora. The wetlands were probably sur- rounded by somewhat unstable dry ground. where 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 t o 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 remains that have been carried out. In addition, nearly all records are the oldest known from 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 considering that pre-Holocene Quaternary deposits are almost unknown from this island (Landvik et al. 1995). The data also show that the present flora, and the range of habitats and plant communities of Svalbard, were already well estab- lished in the earliest Holocene. We are inclined to follow Nathorst’s view that only few vascular plants survived the Weichselian glacial maximum on Svalbard because most lowland areas were glaciated (Mangerud et al. 1992) and because the climate was colder than at present. If this is the case. the major part of the flora must have immi- grated in the latest part of the Weichselian. Many radiocarbon dates between 13 and 10ka from western Spitsbergen show that some areas were deglaciated by then (Mangrud et al. 1992). Deglaciation of lowland areas on Edgeoya began about 10.3 ka BP (Landvik et al. 1995). It should also be pointed out that virtually no clear endemic species occur on Svalbard. a fact which also tes- tifies to the young age of the flora. Acknowledgements. we thank C . Kronborg and 0. Stubdrup. University of Aarhus and P . 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