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1 Disjunctive Distribution of Vaccinium vitis-idaea and Thermal Condition* Woo-seok, Kong** Jong-Hwan, Lim*** 350m 1,200m 6.6 Abstract Vaccinium vitis-idaea or lingonberry, a typical arctic-alpine plant, is common on the circumpolar regions and alpine belts of the Northern Hemisphere, and also occurs on the alpine and subalpine belts of the Korean Peninsula, including Mt. Sorak and Hongchon. Vaccinium vitis-idaea at the elevation of c. 350m a.s.l. of Hongchon is found on the wind hole area with cool summer, and mild winter. Vaccinium vitis-idaea at Hongchon is regarded as the glacial relict of the Pleistocene period, and shows a disjunctive distribution, along with the alpine and subalpine belts of the northern and central Korea since the Holocene period. Present vertical range of Vaccinium vitis-idaea between Mt. Sorak and Hongchon might indicates that the temperatures during the glacial epoch was colder than today, down to -6 to -7 C Vaccinium vitis-idaea at this fragile wind hole site could be endanger if current global warming trends continues, and anthropogenic activities become serious. : Vaccinium vitis-idaea, arctic-alpine plant, Mt. Sorak, Hongchon, disjunctive distribution, wind hole, glacial relict, global warming 2007 (This work was supported by the research grant of the Kyunghee University in 2007). (Professor, Department of Geography and Institute of Basic Sciences, Kyunghee University), wskong@khu.ac.kr (Senior Researcher, Department of Forest Environment, Korea Forest Research Institute), limjh@forest.go.kr 495

2 4 500 relict species endemic species 4 Pleistocene refugia Vaccinium vitisidaea L. 350m ecojournal.co.kr wind hole natural history 200m cm 2 1 2m 3 3 SP-2000 Veriteq Co., Canada US FS 2008a usda.gov USDA ARS 2008 ZipcodeZoo 2008 Vaccinium vitis-idaea L. Kingdom Plantae Subkingdom Tracheobionta Superdivision Spermatophyta Division Magnoliophyta Class Magnoliopsida Dilleniidae Subclass Dilleniidae Order Ericales Family Ericaceae Vaccinioideae Subfamily Vaccinioideae Genus Vaccinium Vitis-Idaea Section Vitis-Idaea Vaccinium vitis-idaea L. 496

3 Dilleniidae 1 Vaccinium Ericaceae lingonberry, cowberry, foxberry, mountain cranberry, red whortleberry, lowbush cranberry, partridgeberry, redberry Oxycoccus Vaccinium subgenera 20 sections 2 Vitisidaea Vaccinium uliginosum Vaccinium go.kr Vaccinium Oxycoccus Vaccinium Oxycoccus, Oxycoccoides Batodendron, Brachyceratium, Bracteata, Ciliata, Cinctosandra, Conchophyllum, Cyanococcus, Eococcus, Epigynium, Galeopetalum, Hemimyrtillus, Myrtillus, Neurodesia, Oarianthe, Oreades, Pachyanthum, Polycodium, Pyxothamnus, Vaccinium, Vitis-idaea Vaccinium bracteatum Vaccinium hirtum var. koreanum Vaccinium japonicum Vaccinium japonicum var. ciliaris Vaccinium oldhamii Vaccinium oldhamii for. glaucinum Vaccinium oxycoccus Vaccinium oxycoccus subsp. microcarpus Vaccinium uliginosum Vaccinium uliginosum for. depressum Vaccinium uliginosum for. ellipticum Vaccinium uliginosum var. alpinum Vaccinium vitis-idaea 497

uliginosum var. alpinum Vaccinium vitis-idaea var. genuinum 8 Hugeria japonica Oxycoccus microparpus Oxycoccus quadripetalus 1979 1997 5 1996 Vaccinium microcarpum 10 2001 Vaccinium japonicum var.

4 uliginosum var. alpinum Vaccinium vitis-idaea var. genuinum 8 Hugeria japonica Oxycoccus microparpus Oxycoccus quadripetalus Vaccinium microcarpum Vaccinium japonicum var. japonicum Vaccinium japonicum var. ciliare Vaccinium koreanum Vaccinium oldhamii var. oldhamii 8 Flora of Korea Editorial Committee Vaccinium Vaccinium uliginosum var. alpinum Vaccinium koreanum 7 20cm 1 3cm 5 13mm mm mm hermaphrodite self-fertile Davis et al Baskin et al Hill & Vander Kloet 2005 Eriksson & Froborg Hall & Shay 1981 BWCA 2008 US FS, 2008b Ritchie 1955 Nekola

5 Henry 2003 Ritchie 1955 Kardell 1986 Rook 2002 Holloway Kuchko Jeglum 1971 Hall & Shay 1981 Holloway 1981 US FS 2008b Fernquist 1977 Launert 1981 PFAF 2008 C A B B1 B2 B3 phytochemical BWCA 2008 Stang et al, Tufts circumpolar circumboreal 2 Hulten

6 Ritchie 1955 Hultén Ohwi 1965 Holloway 1981 USDA ARS 2008 ZipcodeZoo 2008 Hultén 1970 Vaccinium vitis-idaea subsp. vitis-idaea 10 25mm 2 000m Vaccinium vitis-idaea subsp. minus 7 20mm m Fernald 1970 Hultén 1970 subsp. vitis-idaea subsp. minus Kurilo 3 Pliocene 2 Tirmenstein m efloras m 2 700m 2 500m 1 200m Hultén m Landolt m Kolbek and Kucera m m http: //ipcp.edunet4u.net m 1 214m 350m m m

7 1 746m m m m m 894m m m 1 400m m m m m m m 1 600m m a b 1989a b c 1 708m 1 950m 1 915m 10 Pinus pumila Taxus caespitosa Juniperus chinensis var. sargentii Thuja koraiensis Cypripedium guttatum var. koreanum Arctous ruber 501

Silene oliganthella Anemone narcissiflora Pedicularis manshurica Bupleurum euphorbioides Anemone narcissiflora Rosa koreana Rosa marretii Trientalis europaea Lilium

8 Silene oliganthella Anemone narcissiflora Pedicularis manshurica Bupleurum euphorbioides Anemone narcissiflora Rosa koreana Rosa marretii Trientalis europaea Lilium cernuum Iris uniflora var. carina m Vaccinium koreana m 1 676m 1 578m m 1 103m 1 214m 350m talus wind hole Lee & Kim

Cystopteris fragalis Athyrium crenatum Polypodium virginianum Hackelia deflexa 2006 350m 70m 30m 25m 7m 200m 2 5 Sorbaria sorbifolia var.

9 Cystopteris fragalis Athyrium crenatum Polypodium virginianum Hackelia deflexa m 70m 30m 25m 7m 200m 2 5 Sorbaria sorbifolia var. stellipila Betula schmidtii Prunus padus Rhdododendron mucronulatum Diplazium sibiricum Rosa suavis Woodsia subcordata Clematis fusca var. coreana Calamagrostis langsdorfii Carex vaginata var. petersii Dryopteris fragrans Polytrichum commune Iokawa & Ishizawa 2003 Nekola 1999 Iokawa & Ishizawa 2003 palaeorefugia 350m m 1 200m m adiabatic lapse rate mm mm mm mm mm 27mm 503

10 cm m cm Kong & Watts m cm AWS 4 1 2m m cm m cm m m cm 3 1 2m AWS 504

11 m cm Raatikainen & Vanninen 1988 PECC mm m m 350m 350m m 1 200m m 505

12 cm 1 2m cm Ritchie C Wikipedia 2008 Hall & Shay 1981 US FS 2008a Shevtsova et al Holloway 1981 in situ 506

13 core zone buffer zone m 1 816m m

14 a a b c Baskin, C. C., Milberg, P., Andersson, L., and Baskin, J. M., 2000, Germination studies of three dwarf shrubs(vaccinium, Ericaceae) of Northern Hemisphere coniferous forests, Canadian. Journal of Botany, 78(12), Davis, A. N., Holloway, P., and Kruse, J., 2003, Insect visitors and potential pollinators of lingonberries, Vaccinium vitis-idaea subsp. minus in sub-arctic Alaska, Acta Horticulturae, 626, 441~446. Eriksson, O. and Froborg, H., 1996, Windows of opportunity for recruitment in long-lived clonal plants: experimental studies of seedling establishment in Vaccinium shrubs, Canadian. Journal of Botany, 74, Fernald, M. L., 1970, Gray s Manual of Botany, 8th ed., D. Van Nostrand Co., New York. Fernquist, I., 1977, Results of experiments with cowberries and blueberries in Sweden, Acta Horticulturae., 61, 295~300. Flora of Korea Editorial Committee, 2007, The Genera of Vascular Plants of Korea, Academy Publ. Co. Hall, I. V. and J. M. Shay, 1981, The biological flora of Canada, Vaccinium vitis-idaea L. var. minus Lodd., Supplementary Account, Canadian Field-Naturalist, 95(4), 434~464. Henry, C., 2003, Refuge for the Ice Age survivor, Endangered Species Bulletin, 28, 24~26. Hill, N. M. & Vander Kloet, S. P., 2005, Longevity of experimentally buried seed in Vaccinium: relationship to climate, reproductive factors and natural seed banks, Journal of Ecology, 93, Holloway, P. S., 1981, Studies on vegetative and reproductive growth of lingonberry, Vaccinium vitis-idaea L., (Ph, D. Thesis.) Saint Paul, MN: 508

15 University of Minnesota. Holloway, P. S., Van Veldhuizen, R. M., Stushnoff, C., and Wildung, D. W., 1982, Vegetative growth and nutrient levels of lingonberries grown in four Alaskan substrates, Canadian. Journal of Plant Science., 62(4), 969~977. Hultén, E., 1958, The Amphi-Atlantic Plants and Their Phytogeographical Connections, Almqvist and Wiksell, Stockholm. Hultén, E., 1962, The Circumpolar Plants. I, Almqvist and Wiksell, Stockholm. Hultén, E., 1970, The Circumpolar Plants. II, Almqvist and Wiksell, Stockholm. Iokawa, Y. and Ishizawa, S., 2003, Vascular plants of wind-hole areas in Japan (1), J. Journal of Phytogeography and Taxon, 51, 13~26. Jeglum, J. K., 1971, Plant indicators of ph and water level in peatlands at Candle Lake, Saskatchewan, Canadian. Journal of Botany, 49, 1661~1676. Kardell, L., 1986, Occurrence and berry production of Rubus chamaemorus L., Vaccinium oxycoccus L., V. microcarpum Turcz. & V. vitis-idaea on Swedish peatlds, Scandinavian Journal of Forest Research, 1(1), 125~140. Kolbek, J. and Kucera, M., 1989, A Brief Survey of Selected Woody Species on North Korea (D.P.R.K), Botanical Institute, Czechoslovak Academy of Sciences, Czechoslovakia. Kong, W. S. and Watts, D., 1993, The Plant Geography of Korea, Kluwer Publ., The Netherlands. Kuchko, A. A., 1988, Bilberry and cowberry yields and the factors controlling them in the forests of Karelia, U.S.S.R., Acta Botanica, Fennica., 136, 23~25. Landolt, E., 1996, Vaccinium vitis-idaea subsp. minus, An overlooked circumpolar-arctic taxa of the Alps, Anales Jardine de Botanique, Madrid, 54, 277~284. Launert, E., 1981, Edible and Medicinal Plants, Hamlyn. Lee, K. S. and Kim, J. H., 1994, Primary succession on talus area at Mt. Kariwangsan, Korea, Korea Journal of Ecology, 21, 383~387. Nekola, J. C., 1999, Palaerefugia and neorefugia: the influence of colonization history on community pattern and process, Ecology, 80, 2459~2473. Ohwi, J., 1965, Flora of Japan, Washington, DC: Smithsonian Institution. Raatikainen, M. and Vanninen, I., 1988, The effects of the 1984~1985 cold winter on the bilberry and ligonberry yield in Finland, Acta Botanica Fennica, 136, 43~47. Ritchie, J. C., 1955, Vaccinium vitis-idaea, Journal of Ecology, 43(2), 701~708. Rook, E., 2002, Plants of the North, org/earl/bwca/nature/flora.html. Shevtsova, A., Haukioja, E., and Ojala, A., 1997, Growth response of subarctic dwarf shrubs, Empetrum nigrum and Vaccinium vitis-idaea, to manipulated environmental conditions and species removal, Oikos, 78(3), 440~458. Stang, E. J., Weis, G. G., and Klueh, J., 1990, Lingonberry: Potential new fruit for the northern United States, pp. 321~323. In: J. Janick and J. E. Simon (eds.), Advances in New Crops, Timber Press, Portland, OR. Tirmenstein, D., 1991, Vaccinium vitis-idaea, In: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (2003, April). Fire Effects Information System. BWCA (Boundary Water Canoe Area), 2008, The natural history of the northwoods, /earl/bwca/nature/shrubs/vacciniumvit.html (accessed Aug. 2008). efloras. 2008, Flora of China, florataxon.aspx?flora_id=2&taxon_id= (accessed Aug. 2008). PECC, 2008, Plant Ecophysiology and Climate Change Group, University of Helsinki, helsinki.fi/bioscience/pecc (accessed Aug. 2008). PFAF (Plants For A Future), 2008, (accessed Aug. 2008). 509

16 US FS (United States Forest Service), 2008a, Fire Effects Information System, database/feis/plants/shrub/vacvit/all.html (accessed Aug. 2008). US FS(United States Forest Service), 2008b, Fire Effects Information System(FIES), database/feis/plants/shrub/vacvit/all.html#dist RIBUTION%20AND%20OCCURRENCE (accessed Aug. 2008). USDA ARS(United States Department of Agriculture, Agricultural Research Service), 2008, Germplasm Resources Information Network(GRIN), pl?41069 (accessed Aug. 2008). Wikipedia, 2008, (accessed Aug. 2008). ZipcodeZoo, 2008, Vaccinium_vitis-idaea.asp (accessed Aug. 2008) , wskong@khu.ac.kr Correspondence: Woo-seok, Kong, Department of Geography, Kyunghee University, Hoegi-dong, Dongdaemun-gu, , Seoul, Korea(Phone: , wskong@khu.ac.kr) 510

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